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	<front>
		<journal-meta>
			<journal-id journal-id-type="publisher-id">EGEOL</journal-id>
			<journal-title-group>
				<journal-title>Estudios Geol&#xf3;gicos</journal-title>
				<abbrev-journal-title abbrev-type="publisher">Estud. geol.</abbrev-journal-title>
			</journal-title-group>
			<issn publication-format="print">0367-0449</issn>
			<issn publication-format="electronic">1988-3250</issn>
			<issn-l>0367-0449</issn-l>
			<publisher>
				<publisher-name>Consejo Superior de Investigaciones Cient&#xed;ficas</publisher-name>
			</publisher>
		</journal-meta>
		<article-meta>
			<article-id pub-id-type="publisher-id">egeol.44639.613</article-id>
			<article-id pub-id-type="doi">10.3989/egeol.44639.613</article-id>
			<article-categories>
				<subj-group subj-group-type="heading">
					<subject>Articles</subject>
				</subj-group>
			</article-categories>
			<title-group>
				<article-title>First record of fossil sauropterygians from the Upper Triassic of Southwestern Spain (Ayamonte, Huelva province)</article-title>
				<trans-title-group xml:lang="es">
					<trans-title>Primer registro de sauropterigios f&#xf3;siles del Tri&#xe1;sico Superior del suroeste de Espa&#xf1;a (Ayamonte, provincia de Huelva)</trans-title>
				</trans-title-group>
			</title-group>
			<contrib-group>
				<contrib contrib-type="author" corresp="yes">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-4211-3946</contrib-id>
					<name>
						<surname>Reolid</surname>
						<given-names>Mat&#xed;as</given-names>
					</name>
					<email xlink:href="mreolid@ujaen.es">mreolid@ujaen.es</email>
					<aff id="aff1"><institution content-type="department">Departamento de Geolog&#xed;a</institution>, <institution>Universidad de Ja&#xe9;n</institution>, <addr-line>Ja&#xe9;n</addr-line>, <country>Spain</country>. </aff>
				</contrib>
				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-5727-3646</contrib-id>
					<name>
						<surname>Mu&#xf1;iz</surname>
						<given-names>Fernando</given-names>
					</name>
					<aff id="aff2"><institution content-type="department">Departamento de Cristalograf&#xed;a, Mineralog&#xed;a y Qu&#xed;mica Agr&#xed;cola</institution>, <institution>Universidad de Sevilla</institution>, <addr-line>Sevilla</addr-line>, <country>Spain</country>.</aff>
				</contrib>
				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-2144-5714</contrib-id>
					<name>
						<surname>Toscano</surname>
						<given-names>Antonio</given-names>
					</name>
					<aff id="aff3"><institution content-type="department">Departamento de Ciencias de la Tierra</institution>, <institution>Universidad de Huelva</institution>, <addr-line>Huelva</addr-line>, <country>Spain</country>.</aff>
				</contrib>
				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-1707-9670</contrib-id>
					<name>
						<surname>Bela&#xfa;stegui</surname>
						<given-names>Zain</given-names>
					</name>
					<aff id="aff4"><institution content-type="department">Departament de Din&#xe0;mica de la Terra i de l&#x2019;Oce&#xe0;</institution>, <institution content-type="faculty">Facultat de Ci&#xe8;ncies de la Terra</institution>, <institution>Universitat de Barcelona</institution>, <addr-line>Barcelona</addr-line>, <country>Spain</country></aff>
					<aff id="aff5"><institution content-type="institute">Institut de Recerca de la Biodiversitat (IRBio)</institution>, <institution>Universitat de Barcelona</institution>, <addr-line>Barcelona</addr-line>, <country>Spain</country></aff>
				</contrib>
			</contrib-group>
			<pub-date pub-type="epub">
				<day>07</day>
				<month>07</month>
				<year>2022</year>
			</pub-date>
			<pub-date pub-type="collection">
				<month>06</month>
				<year>2022</year>
			</pub-date>
			<volume>78</volume>
			<issue>1</issue>
			<elocation-id>e145</elocation-id>
			<history>
				<date date-type="received">
					<day>19</day>
					<month>03</month>
					<year>2022</year>
				</date>
				<date date-type="accepted">
					<day>09</day>
					<month>06</month>
					<year>2022</year>
				</date>
				<date date-type="rev-recd">
					<day>30</day>
					<month>06</month>
					<year>2022</year>
				</date>
				<date date-type="pub">
					<day>20</day>
					<month>07</month>
					<year>2022</year>
				</date>
			</history>
			<permissions>
				<copyright-statement>&#xa9; 2022 CSIC</copyright-statement>
				<copyright-year>2022</copyright-year>
				<license license-type="open-access" xlink:href="https://creativecommons.org/licenses/by-nc/4.0/">
					<license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution- Non Commercial (by-nc) Spain 4.0 License.</license-p>
				</license>
			</permissions>
			<self-uri xlink:href="http://estudiosgeol.revistas.csic.es/index.php/estudiosgeol/article/view/XXXX/XXXX"/>
			<abstract>
				<title>Abstract</title>
				<p>This work reports the first record of a sauropterygian reptile remain from the uppermost Triassic of the westernmost part of the South Iberian Palaeomargin. The fossil bone, found in the Upper Triassic carbonate succession of Ayamonte (Huelva, Spain), corresponds to a neural arch of a sauropterygian. The carbonate succession was deposited in very shallow marine environment simultaneous with igneous activity during the Rhaetian (latest Triassic). The studied remain is isolated, disarticulated and presents fractures that evidence transport but also potential activity of scavengers. The neural arch is preserved as calcium phosphate enriched in some elements (e.g. Sr) relative to the surrounding carbonate sediment. The anatomic features do not allow a detailed taxonomic identification. This remain extends the record of sauropterygians to the westernmost end of the Tethys following the South Iberian Palaeomargin and evidences the colonization of the Algarve Basin during the extensional phase related with the progress of the rifting of Pangaea and the opening of the Tethys to the west.</p>
			</abstract>
			<trans-abstract xml:lang="es">
				<title>Resumen</title>
				<p>Este trabajo se centra en el estudio del primer registro de un resto f&#xf3;sil de sauropterigio procedente del Tri&#xe1;sico superior del sector oriental del Paleomargen Surib&#xe9;rico. El resto, encontrado en la sucesi&#xf3;n sedimentaria carbonatada del Tri&#xe1;sico que aflora en Ayamonte (Huelva, Espa&#xf1;a), corresponde a un arco neural de un sauropterigio. La sucesi&#xf3;n carbonatada se deposit&#xf3; en un ambiente marino muy somero simult&#xe1;neamente a cierta actividad &#xed;gnea durante el Rhaetiense (Tri&#xe1;sico superior). El resto f&#xf3;sil aparece aislado, desarticulado y presenta algunas fracturas que evidencian cierto transporte por corrientes, sin descartar la posible interacci&#xf3;n de organismos carro&#xf1;eros. El arco neural est&#xe1; conservado como fosfato c&#xe1;lcico con enriquecimiento en Sr, de acuerdo con los mapeos composicionales realizados sobre el resto f&#xf3;sil y el sedimento circundante. Los rasgos anat&#xf3;micos no han permitido su identificaci&#xf3;n taxon&#xf3;mica detallada. El resto estudiado extiende el registro de reptiles sauropterigios hacia el occidente colonizando el Paleomargen Surib&#xe9;rico, en este caso la Cuenca del Algarve, conforme se produc&#xed;a la rotura (<italic>rifting</italic>) de Pangea y el Tethys se abr&#xed;a paso hacia el oeste con la progresiva inundaci&#xf3;n de nuevas cuencas.</p>
			</trans-abstract>
			<kwd-group>
				<kwd>Fossil vertebrate</kwd>
				<kwd>Germanic facies</kwd>
				<kwd>Upper Triassic</kwd>
				<kwd>Westernmost Tethys</kwd>
			</kwd-group>
			<kwd-group xml:lang="es">
				<kwd>Vertebrado f&#xf3;sil</kwd>
				<kwd>Facies Germ&#xe1;nicas</kwd>
				<kwd>Tri&#xe1;sico superior</kwd>
				<kwd>Tethys Occidental</kwd>
			</kwd-group>
			<funding-group id="fw-01">
				<award-group id="aw1">
					<funding-source>Junta de Andaluc&#xed;a</funding-source>
					<award-id>PY20_00111</award-id>
					<award-id>RNM-200</award-id>
				</award-group>
				<award-group id="aw2">
					<funding-source>Spanish Govern</funding-source>
					<award-id>PID2019-104625RB-100</award-id>
				</award-group>
				<funding-statement>Authors would like to thank Ram&#xf3;n Mart&#xed;n, Miguel &#xc1;ngel Bernal and Benjam&#xed;n Cabaco for informing us of the discovery of the fossil bone. Financial support through the projects PY20_00111 and RNM-200 Research Group (Junta de Andaluc&#xed;a, Spain) and PID2019-104625RB-100 (Spanish Govern) is gratefully acknowledged. Technical and human support provided by Centro de Instrumentaci&#xf3;n Cient&#xed;fico-T&#xe9;cnica (CICT) of the University of Ja&#xe9;n is gratefully acknowledged. The authors thank the constructive comments of the reviewers Carlos de Miguel Chaves and Torsten Scheyer.</funding-statement>
			</funding-group>
			<counts>
				<fig-count count="3"/>
				<table-count count="0"/>
				<equation-count count="0"/>
				<ref-count count="57"/>
				<page-count count="10"/>
			</counts>
		</article-meta>
	</front>
	<body>
		<sec id="sec1" sec-type="intro">
			<title>Introduction</title>
			<p>Marine reptiles were diverse and abundant in the Triassic marine ecosystems of Western Tethys, both in the Germanic and in the Alpine basins, forming part of the Germanic Bioprovince and the Tethyan Bioprovince, as compiled in <xref ref-type="bibr" rid="B39">Rieppel (2000)</xref> and <xref ref-type="bibr" rid="B27">Motani (2009)</xref>. In the Alpine Triassic, in addition to Sauropterygia, Ichthyosauria and Thalattosauria have also been recorded (e.g. <xref ref-type="bibr" rid="B28">M&#xfc;ller, 2005</xref>; <xref ref-type="bibr" rid="B9">Dalla Vecchia, 2006</xref>, <xref ref-type="bibr" rid="B10">2008</xref>; <xref ref-type="bibr" rid="B18">Kolb <italic>et al</italic>., 2011</xref>). The Besano Formation in the Southern Alps from Italy and Switzerland, is especially rich in marine reptiles (e.g. <xref ref-type="bibr" rid="B51">Tschanz, 1989</xref>; <xref ref-type="bibr" rid="B3">Beardmore &amp; Furrer, 2016</xref>; <xref ref-type="bibr" rid="B40">Rieppel, 2019</xref>). At the Germanic Basin, from the Muschelkalk facies, the most abundant reptiles in shallow marine sediments were Sauropterygia (e.g. <xref ref-type="bibr" rid="B39">Rieppel, 2000</xref>; <xref ref-type="bibr" rid="B17">Klein <italic>et al</italic>., 2015</xref>). </p>
			<p>The Germanic facies also developed along the Iberian Palaeomargin, today outcropping in the Algarve Basin, the Betic External Zones, the Iberian Range, the Catalonian Coastal Range and the Pyrenean Cordillera. The record of marine reptiles in these areas is also relatively rich but most of the reported remains consist of isolated elements (e.g. <xref ref-type="bibr" rid="B45">Sanz, 1976</xref>, <xref ref-type="bibr" rid="B46">1983a</xref>, <xref ref-type="bibr" rid="B47">b</xref>; <xref ref-type="bibr" rid="B1">Alafont, 1992</xref>; <xref ref-type="bibr" rid="B48">Sanz <italic>et al</italic>., 1993</xref>; <xref ref-type="bibr" rid="B30">Niemeyer, 2002</xref>; <xref ref-type="bibr" rid="B34">Quesada &amp; Aguera Gonz&#xe1;lez, 2005</xref>; <xref ref-type="bibr" rid="B14">Fortuny <italic>et al</italic>., 2011</xref>; <xref ref-type="bibr" rid="B35">Reolid <italic>et al</italic>., 2014</xref>; <xref ref-type="bibr" rid="B11">de Miguel Chaves <italic>et al.</italic>, 2015</xref>, <xref ref-type="bibr" rid="B23">2016</xref>, <xref ref-type="bibr" rid="B24">2017</xref>, <xref ref-type="bibr" rid="B25">2018</xref>, <xref ref-type="bibr" rid="B26">2020</xref>; <xref ref-type="bibr" rid="B26">Campos &amp; Mateus, 2018</xref>; <xref ref-type="bibr" rid="B21">M&#xe1;rquez-Aliaga <italic>et al</italic>., 2019</xref>; <xref ref-type="bibr" rid="B32">P&#xe9;rez-Valera <italic>et al</italic>., 2020</xref>; <xref ref-type="bibr" rid="B43">Ruci&#x144;ski, 2020</xref>). Only one record of sauropterygians in the Iberian Peninsula comes from Alpine facies, being an isolated rib from the Ladinian of the Alpujarride Complex (Betic Internal Zones; <xref ref-type="bibr" rid="B36">Reolid &amp; Reolid, 2020</xref>). </p>
			<p>In the southwestern part of the Iberian Palaeomargin the record of marine reptiles is comparatively scarce. Only at the Algarve Basin (South Portugal), in the Carnian Gr&#xe9;s de Silves Group, fossil vertebrates have been reported from the Rocha da Pena bonebed with numerous temnospondyl remains of <italic>Metoposaurus algarvensis</italic> (<xref ref-type="bibr" rid="B5">Brusatte <italic>et al</italic>., 2015</xref>). In the Gr&#xe9;s de Silves Group have been also reported a probably basal representative of phytosaurs (<xref ref-type="bibr" rid="B22">Mateus <italic>et al</italic>., 2014</xref>) and the placodont <italic>Henodus</italic>; the latter represented by isolated remains including osteoderms and cranial remains (<xref ref-type="bibr" rid="B7">Campos <italic>et al</italic>., 2017</xref>; <xref ref-type="bibr" rid="B6">Campos &amp; Mateus, 2018</xref>; <xref ref-type="bibr" rid="B43">Ruci&#x144;ski, 2020</xref>). </p>
			<p>The objective of this short note is reporting the first record of an isolated vertebra of sauropterygians from the Upper Triassic of the eastern part of the Algarve Basin in Ayamonte (Huelva province, Southwestern Spain).</p>
		</sec>
		<sec id="sec2">
			<title>Geological setting</title>
			<p>The studied remain was recorded in the Triassic deposits that outcrops in the east side of the Guadiana River, close to Ayamonte town, and more exactly in the hill of the Parador Nacional (coord. 37&#xba;13&#xb4;32&#xb4;&#xb4;N, 7&#xba;24&#xb4;26&#xb4;&#xb4;W) (<xref ref-type="fig" rid="f1">Fig. 1A</xref> and <xref ref-type="fig" rid="f1">B</xref>). The Triassic of Ayamonte constitute the eastern end of the Algarve Basin. This basin is extended in E-W direction from Ayamonte (Spain) to Cape San Vicente (Portugal) and constitutes the westernmost part of the South Iberian Palaeomargin.</p>
			<fig id="f1">
				<label>Figure 1</label>
				<caption>
					<title>Geological setting.</title>
					<p>A. Geological map of southwestern Iberia with indication of Ayamonte Town. B. Geological map of the surroundings of the Ayamonte (Huelva) with location of the outcrop with fossil remain (modified from <xref ref-type="bibr" rid="B2">Alonso-Chaves et al., 2020</xref>). C. Lithological columns on the eastern margin of the Guadiana River (modified from <xref ref-type="bibr" rid="B2">Alonso-Chaves et al., 2020</xref>) and location of the fossil bone.</p>
				</caption>
				<graphic id="gra-1" xlink:href="EGEOL-78-01-e145-gf1.png"/>
			</fig>
			<p>In Ayamonte, the Triassic is unconformably overlying the shales of the Lower Carboniferous. The lower part of the Triassic succession (around 15 m thick) is constituted by red siltstones and sandstones with gypsum and some microconglomerate intervals. These deposits were interpreted as related to continental or coastal deposits (<xref ref-type="bibr" rid="B2">Alonso-Chaves <italic>et al</italic>., 2020</xref>; <xref ref-type="bibr" rid="B44">Santos <italic>et al</italic>., 2022</xref>). The trace fossils (<italic>Taenidium</italic> isp. <italic>Labyrintichnus terrerensis</italic>, <italic>Planolites</italic> isp.) are indicative of the <italic>Scoyenia</italic> ichnofacies typical of swamp-like to alluvial plain environments (<xref ref-type="bibr" rid="B44">Santos <italic>et al</italic>., 2022</xref>). These deposits are the continuation (in the eastern side of the Guadiana River) of the Gr&#xe9;s de Silves Group, which was described in the Portuguese part of the Algarve Basin and assigned to the Carnian by <xref ref-type="bibr" rid="B31">Palain (1976)</xref>. The beginning of the sedimentation in this basin occurred during the early Carnian according to the palynological analysis of <xref ref-type="bibr" rid="B54">Vilas-Boas <italic>et al</italic>. (2022)</xref>. </p>
			<p>The upper part of the Triassic sedimentary succession (around 26 m thick) is constituted by a carbonate interval composed by marlstones, marly-limestones, limestones and dolostones interbedded with basic volcanic rocks (dolerites) and volcano-sedimentary deposits to the top (see detailed description in <xref ref-type="bibr" rid="B2">Alonso-Chaves <italic>et al</italic>., 2020</xref>; <xref ref-type="fig" rid="f1">Fig. 1C</xref>). The faunal assemblage of the succession is dominated by bivalves (mainly <italic>Trigonodus</italic>, <italic>Isocyprina</italic>, <italic>Pleuromya</italic> and <italic>Isognonom</italic>) and gastropods (mainly <italic>Coelostylina</italic>, <italic>Spirostylus</italic> and <italic>Mathilda</italic>). <xref ref-type="bibr" rid="B44">Santos <italic>et al</italic>. (2022)</xref> recorded <italic>Palaeonucula subovalis</italic>, <italic>Modiolus</italic> cf. <italic>minimus</italic>, and <italic>Isocyprina concentrica</italic>; bivalves indicative of a Rhaetian age (<xref ref-type="bibr" rid="B55">V&#xf6;r&#xf6;s, 1981</xref>; <xref ref-type="bibr" rid="B20">M&#xe1;rquez-Aliaga <italic>et al</italic>., 2010</xref>). The stratigraphic record of other genera of mollusks reported from this section support the assignation to the uppermost Triassic. According to <xref ref-type="bibr" rid="B42">Ros (2009)</xref> and <xref ref-type="bibr" rid="B20">M&#xe1;rquez-Aliaga <italic>et al</italic>. (2010)</xref>, the first occurrence of bivalves <italic>Pteromya</italic> cf. <italic>tatei</italic> and the genus <italic>Isocyprina</italic> is in Rhaetian rocks. The first occurrence of the gastropod <italic>Cylindrobullina</italic> cf. <italic>avenoides</italic> in Europe and South America is also recorded in the Rhaetian (<xref ref-type="bibr" rid="B13">Ferrari, 2015</xref>). </p>
			<p>According to the presence of trace fossils (<italic>Thalassinoides</italic> isp., <italic>Treptichnus pollardi</italic> and <italic>Helminthoidichnites tenuis</italic>), fossil macroinvertebrates and sedimentary structures, <xref ref-type="bibr" rid="B44">Santos <italic>et al</italic>. (2022)</xref> have interpreted a shallow marginal environment with brackish and low energy conditions.</p>
		</sec>
		<sec id="sec3" sec-type="materials|methods">
			<title>Materials and methods</title>
			<p>The studied remain appeared in the broken surface of a red limestone block and it is constituted by two halves identified as NOT/AY/001a and NOT/AY/001b. Polished slabs and thin sections of the red limestone block have been prepared and the microfacies being analysed with an Olympus SZ60 microscope at the Universidad de Ja&#xe9;n.</p>
			<p>The fossil bone and hosting rock were scanned at the Universidad de Ja&#xe9;n using a Bruker XR-microfluorescence M4 Tornado equipped with a rhodium target X-ray tube with a high voltage of 50 kV, a current of 600 &#x3bc;A and pressure of 20 mbar. The spotsize of the X-ray optics was 25 &#x3bc;m. The maximum penetration depth from which fluorescence X-rays can still reach the detector is less than 20 &#x3bc;m. This low penetration allowed analyses of the sediment surface showing lateral compositional changes, especially the contrast between the fossil bone and the surrounding sedimentary rock. The geochemical compositional maps obtained for each element are represented by a range of colour intensity that indicates the relative concentration of each element.</p>
			<sec id="sec3.1">
				<title>Results</title>
				<p>The fossil bone is recorded in red to purple limestone which is roughly laminated with calcisiltite to calcarenite thin layers, locally rich in organic matter and native sulfur. The sediment surrounding the fossil bone is constituted by a bioclastic packstone of peloids, lumps and thin-shelled bivalves. Bivalves are commonly disarticulated and concave-up in the bed, most of them &lt; 1 cm (<xref ref-type="fig" rid="f2">Fig. 2A</xref>). Microgastropods and coal fragments are also common in the bed with the fossil bone.</p>
				<fig id="f2">
					<label>Figure 2</label>
					<caption>
						<title>A. Polished slab showing the presence of abundant thin-shelled disarticulated bivalves. B. View of the neural arch of sauropterygian. C. Sketch of the neural arch with indication of anatomic parts. D. Detail of the spongy tissue preserved in the fossil bone with infilling of iron oxides and growth of pyrolusite in the bone surface.</title>
					</caption>
					<graphic id="gra-2" xlink:href="EGEOL-78-01-e145-gf2.png"/>
				</fig>
				<p>The fossil bone is an isolated vertebral arch without centrum (<xref ref-type="fig" rid="f2">Figs. 2b</xref> and <xref ref-type="fig" rid="f2">2C</xref>). The maximum width of the neural arch is 46 mm and the maximum height is 44 mm. The neural arch presents a high and thin neural spine (31 mm high and 4 mm width). The square-sided transverse processes are thick (10 mm high and 21 mm width) and the distal margins are slightly rounded. The transverse processes present areas with spongy bone tissue. The neural canal or vertebral foramen is elevated and relatively circular with 8 mm in diameter. The development of the transverse apophysis sensu <xref ref-type="bibr" rid="B1">Alafont (1992)</xref> (maximum width of neural arch / height of neural canal) is 5.76, whereas the development of the neural canal (neural canal height*100 / neural arch height) is 18.26. Since the studied specimen is a section in a fracture surface, it is not possible to study the morphology of the zygosphene-zygantrum articulation. </p>
				<p>The upper part of the left transverse process presents a semicircular fracture with around 1 cm in diameter (<xref ref-type="fig" rid="f2">Fig. 2B</xref> and <xref ref-type="fig" rid="f2">C</xref>). Other minor fractures are evidenced in other places of the transverse processes and the neural spine. The spongy bone tissue present infilling by iron oxides whereas pyrolusite is locally observable at the surface (<xref ref-type="fig" rid="f2">Fig. 2D</xref>). </p>
				<p>According to the geochemical compositional maps (<xref ref-type="fig" rid="f3">Fig. 3</xref>), the bone is preserved as calcium phosphate. The Sr content is especially high in the bone and is absent in the surrounding sediment which is enriched in Si, Fe, Ba and Cu in respect to the bone. </p>
				<fig id="f3">
					<label>Figure 3</label>
					<caption>
						<title>Compositional maps of XR-microfluorescence of the fossil neural arch highlighting the distribution of Ca, P, Sr, Fe, Ba and Cu. Note the intensity of colour indicate more relative concentration.</title>
					</caption>
					<graphic id="gra-3" xlink:href="EGEOL-78-01-e145-gf3.png"/>
				</fig>
			</sec>
		</sec>
		<sec id="sec4" sec-type="discussion">
			<title>Discussion</title>
			<sec id="sec4.1">
				<title>Taxonomic attribution</title>
				<p>The isolated neural arch is attributed to a sauropterygian as evidenced by the morphology of the thin and elongated neural spine with relatively reduced development of the square-sided transverse processes. These features allow to exclude the assignation to phytosaurs, because this group exhibit more robust vertebrae with thicker neural spines for supporting dorsal armor plates and thick and large transverse processes (<xref ref-type="bibr" rid="B8">Case, 1932</xref>; <xref ref-type="bibr" rid="B19">Lucas <italic>et al</italic>., 2002</xref>; <xref ref-type="bibr" rid="B57">Witzmann <italic>et al</italic>., 2014</xref>). Other typical aquatic diapsid reptiles of Late Triassic (<xref ref-type="bibr" rid="B29">Nicholls, 1999</xref>; <xref ref-type="bibr" rid="B41">Rieppel <italic>et al</italic>., 2000</xref>), the thalattosaurians, are also excluded due to the characteristic small neural canal of vertebrae in spite of the neural archs of thalattosaurs are of moderate height, almost in the Superfamily Askeptosauroidea (<xref ref-type="bibr" rid="B28">M&#xfc;ller, 2005</xref>). </p>
				<p>The development of the transverse apophysis shows a ratio of 5.76 whereas this value is around 7.3 in the Family Pachypleurosauridae and ranges between 3.7 to more than 8.5 in Placodontia (<xref ref-type="bibr" rid="B1">Alafont, 1992</xref>). Transverse processes of the dorsal vertebrae are specially elongated in placodonts such as <italic>Placodus gigas</italic> (<xref ref-type="bibr" rid="B41">Rieppel, 2000</xref>). This ratio in the dorsal vertebrae documented by <xref ref-type="bibr" rid="B49">Segesdi &amp; Osi (2021)</xref> ranges from 5.2 to 6.4 for <italic>Nothosaurus</italic> and is around 5 for simosaurids. </p>
				<p>The development of the neural canal in the studied vertebra is 18.26, whereas this ratio is 27.8 in pachypleurosaurids and from 25 to 31 in placodontids (<xref ref-type="bibr" rid="B1">Alafont, 1992</xref>). According to the vertebrae reported by <xref ref-type="bibr" rid="B49">Segesdi &amp; Osi (2021)</xref>, the development of the neural canal is lower in <italic>Nothosaurus</italic> (10.9 to 12.9) and in simosaurids (8.6). However, the values proposed by <xref ref-type="bibr" rid="B1">Alafont (1992)</xref> comparing the development of transverse processes and the development of the neural canal, are variable along the postcranial skeleton from cervical to caudal vertebrae. Therefore, the taxonomic assignation of the vertebra within Sauropterygia is complicated. Pachypleurosaurids can be excluded because their size was much smaller than the studied specimen. In fact, this specimen was relatively large due to the size of the neural arch, probably reaching more than 2 m in length. But an adult specimen of <italic>Nothosaurus</italic> presents larger dorsal vertebrae than the studied specimen, reaching more than 8 cm height for the neural spine and 6 cm width for the neural arch (<xref ref-type="bibr" rid="B49">Segesdi &amp; Osi, 2021</xref>). Therefore, the studied neural arch could correspond to a caudal vertebrae of an adult <italic>Nothosaurus</italic>.</p>
				<p>Nevertheless, considering the age of the studied fossil remain (Upper Triassic, probably Rhaetian), the fossil record of sauropterygians (<xref ref-type="bibr" rid="B38">Rieppel, 1999</xref>), and the features of the neural arch, the studied remain would also correspond to a cyamodontoid placodont such as <italic>Henodus</italic> or <italic>Psephoderma</italic>. Most of the sauropterygians, included other placodonts, nothosauroids and pistosauroids, are mainly restricted to Middle Triassic (see <xref ref-type="bibr" rid="B38">Rieppel, 1999</xref>). <italic>Henodus</italic> was recorded from the Algarve as Carnian (<xref ref-type="bibr" rid="B6">Campos &amp; Mateus, 2018</xref>; <xref ref-type="bibr" rid="B43">Ruci&#x144;ski, 2020</xref>) and only <italic>Psephoderma</italic> has been reported from Rhaetian (<xref ref-type="bibr" rid="B38">Rieppel, 1999</xref>). Recently, a new placodont, <italic>Parahenodus</italic>, has been described from the Upper Triassic (Carnian-Norian) of Spain (de <xref ref-type="bibr" rid="B25">Miguel Chaves <italic>et al</italic>., 2018</xref>). Although this genus has been described from a unique partial skull (vertebrae have not been recorded), it could also be considered as a potential candidate for the taxonomic assignation of the studied remain. If this assignation to Superfamily Cyamodontoidea is accepted, this would be a neural arch from a caudal vertebra and not from a dorsal vertebra of cyamodontoids. The dorsal vertebrae of cyamondontoids present characteristically elongated, broaded and curved transverse processes, and these features are not observed in the studied neural arch of Ayamonte. However, the neural canal of cyamodontoids is relatively high and narrow (<xref ref-type="bibr" rid="B39">Rieppel, 2000</xref>) distinct to the studied specimen and the taxonomic uncertainty persists. </p>
				<p>The characteristic compact vertebrae of sauropterygians are usually well preserved but the studied specimen present numerous fractures. The typically high neural spines of the sauropterygians are commonly well preserved but commonly disarticulated respect to the centrum, as occurs in the studied vertebra. The disarticulation of centra and neural arches is typical in bonebeds (<xref ref-type="bibr" rid="B35">Reolid <italic>et al</italic>., 2014</xref>). The main distortions in the vertebrae are fractures in neural spines and transverse processes as here reported. The fossil vertebra is isolated and other fossil bones have not been recorded, which is an evidence of transport and dispersion of bone remains by currents and potentially scavengers. Fragmentation of the bone points to a more or less extensive bioestratinomic exposure of the bone previous to the definitive burial.</p>
			</sec>
			<sec id="sec4.2">
				<title>Geochemical composition</title>
				<p>The main composition of specimen (calcium phosphate) does not seem to have been modified during diagenesis. However, the observed enrichment in Sr could be interpreted as related to the replacement of Ca in bioapatite (hydroxylapatite) via co-precipitation or adsorption. Sr is commonly incorporated into bones as a biogenic trace element (e.g. <xref ref-type="bibr" rid="B4">Bocherens <italic>et al</italic>., 1994</xref>; <xref ref-type="bibr" rid="B15">Gilbert <italic>et al</italic>., 1994</xref>; <xref ref-type="bibr" rid="B50">Silen &amp; Sealy, 1995</xref>; <xref ref-type="bibr" rid="B16">Keenan <italic>et al</italic>., 2016</xref>; <xref ref-type="bibr" rid="B37">Rey <italic>et al</italic>., 2022</xref>). However, since Ba and Sr are incorporated by living organisms in the same way, and the Ba content in the studied bone is not remarkable (<xref ref-type="fig" rid="f3">Fig. 3</xref>), probably part of the Sr has a diagenetic origin (<xref ref-type="bibr" rid="B52">Tuken <italic>et al</italic>., 2008</xref>). Sr-rich fluids may also have leached from sulfate-rich Triassic deposits or from doleritic volcanic rocks (<xref ref-type="bibr" rid="B56">Wey <italic>et al</italic>., 2022</xref>). Therefore, Sr was very likely incorporated via adsorption in bioapatite during diagenesis. </p>
				<p>On the other hand, the presence of iron and manganese oxides must be related to diagenetic processes. Iron oxides are infilling small voids of the spongy bone tissue and manganese oxides (pyrolusite) are located on the bone surface but not within the bone. The parallel distribution of Fe and Cu in the spongy bone tissue could be related to the presence of original sulphides (<xref ref-type="bibr" rid="B52">Tusken <italic>et al</italic>., 2008</xref>) related to microbial activity associated with decay of the soft tissues (e.g. <xref ref-type="bibr" rid="B53">Vietti <italic>et al</italic>., 2015</xref>; <xref ref-type="bibr" rid="B33">Pesquero <italic>et al</italic>., 2015</xref>; <xref ref-type="bibr" rid="B12">Domenech-Carbo <italic>et al</italic>., 2016</xref>).</p>
			</sec>
			<sec id="sec4.3">
				<title>Sedimentary environment</title>
				<p>Most of the specimens of sauropterygians recorded in Iberia are Middle Triassic in age (see <xref ref-type="bibr" rid="B26">de Miguel Chaves <italic>et al</italic>., 2020</xref>), but this specimen was recovered from the less common Upper Triassic record. In addition, this specimen, together with the placodont remains of the Algarve Basin (<xref ref-type="bibr" rid="B6">Campos &amp; Mateus, 2018</xref>; <xref ref-type="bibr" rid="B43">Ruci&#x144;ski, 2020</xref>), constitute the westernmost record of Triassic sauropterygians in the Tethys. The detrital red beds (silts and sands) of the Silves Group represent the earliest phase of sedimentation related to the initial rifting of Pangaea, dated as early Carnian (<xref ref-type="bibr" rid="B54">Vilas-Boas <italic>et al</italic>., 2022</xref>). The carbonate succession, dated as Rhaetian (<xref ref-type="bibr" rid="B44">Santos <italic>et al</italic>., 2022</xref>), represents the flooding of the Algarve Basin and the colonization by mollusks (bivalves and gastropods) and marine vertebrates such as phytosaurs (<xref ref-type="bibr" rid="B22">Mateus <italic>et al</italic>., 2014</xref>), the placodont <italic>Henodus</italic> (<xref ref-type="bibr" rid="B6">Campos &amp; Mateus, 2018</xref>; <xref ref-type="bibr" rid="B43">Ruci&#x144;ski, 2020</xref>) and the here studied sauropterygian of Ayamonte. The presence of dolerite rocks in the succession is congruent with the extensional phase of the rifting.</p>
			</sec>
		</sec>
		<sec id="sec5" sec-type="conclusions">
			<title>Conclusions</title>
			<p>The studied remain corresponds to a neural arch of an Upper Triassic sauropterygian deposited in a very shallow marine environment. The studied remain is isolated, disarticulated and presents fractures, that evidence transport but also potential activity of scavengers. The neural arch is preserved as calcium phosphate and some elements such as Sr, are especially rich compared with the surrounding carbonate sediment; fact that could probably be related to substitution of Ca by Sr during the diagenesis. Although the anatomic features do not allow a more accurate taxonomic identification, the studied remain probably corresponds to the neural arch of a caudal cyamodontoid vertebra. This remain extends the record of sauropterygians to the westernmost end of the Tethys and evidences the colonization of the Algarve Basin during the extensional phase related with the progress of the rifting of Pangaea and the opening of the Tethys to the west.</p>
		</sec>
	</body>
	<back>
		<ack>
			<title>Acknowledgements</title>
			<p>Authors would like to thank Ram&#xf3;n Mart&#xed;n, Miguel &#xc1;ngel Bernal and Benjam&#xed;n Cabaco for informing us of the discovery of the fossil bone. Financial support through the projects PY20_00111 and RNM-200 Research Group (Junta de Andaluc&#xed;a, Spain) and PID2019-104625RB-100 (Spanish Govern) is gratefully acknowledged. Technical and human support provided by Centro de Instrumentaci&#xf3;n Cient&#xed;fico-T&#xe9;cnica (CICT) of the University of Ja&#xe9;n is gratefully acknowledged. The authors thank the constructive comments of the reviewers Carlos de Miguel Chaves and Torsten Scheyer.</p>
		</ack>
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