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<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Mar. Sci.</journal-id>
<journal-title>Frontiers in Marine Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Mar. Sci.</abbrev-journal-title>
<issn pub-type="epub">2296-7745</issn>
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<publisher-name>Frontiers Media S.A.</publisher-name>
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<article-meta>
<article-id pub-id-type="doi">10.3389/fmars.2024.1347897</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Marine Science</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Ecomorphological adaptation of <italic>Scorpaena porcus</italic> (Linnaeus, 1758): evidence from two different environments revealed by <italic>sagittae</italic> features and somatic growth rates</article-title>
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<contrib contrib-type="author">
<name>
<surname>D&#x2019;Iglio</surname>
<given-names>Claudio</given-names>
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<sup>1</sup>
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<surname>Famulari</surname>
<given-names>Sergio</given-names>
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<xref ref-type="aff" rid="aff1">
<sup>1</sup>
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<name>
<surname>Ferri</surname>
<given-names>Josipa</given-names>
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<xref ref-type="aff" rid="aff2">
<sup>2</sup>
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<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
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<contrib contrib-type="author" corresp="yes">
<name>
<surname>Albano</surname>
<given-names>Marco</given-names>
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<xref ref-type="aff" rid="aff3">
<sup>3</sup>
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<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
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<contrib contrib-type="author">
<name>
<surname>Span&#xf2;</surname>
<given-names>Nunziacarla</given-names>
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<xref ref-type="aff" rid="aff1">
<sup>1</sup>
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<xref ref-type="aff" rid="aff4">
<sup>4</sup>
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<name>
<surname>Capillo</surname>
<given-names>Gioele</given-names>
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<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<xref ref-type="author-notes" rid="fn003">
<sup>&#x2020;</sup>
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<name>
<surname>Savoca</surname>
<given-names>Serena</given-names>
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<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<xref ref-type="author-notes" rid="fn003">
<sup>&#x2020;</sup>
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<aff id="aff1">
<sup>1</sup>
<institution>Department of Chemical, Biological, Pharmaceutical and Environmental Sciences, University of Messina</institution>, <addr-line>Messina</addr-line>, <country>Italy</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Department of Marine Studies, University of Split</institution>, <addr-line>Split</addr-line>, <country>Croatia</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Department of Veterinary Sciences, University of Messina</institution>, <addr-line>Messina</addr-line>, <country>Italy</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Department of Chemical, Biological, Pharmaceutical and Environmental Sciences, Sea in Health and Life Srl, Capo Peloro</institution>, <addr-line>Messina</addr-line>, <country>Italy</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Cristina Porcu, University of Cagliari, Italy</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Andrea Bellodi, University of Cagliari, Italy</p>
<p>Marilena Donnaloia, COISPA Tecnologia &amp; Ricerca, Italy</p>
<p>Jorge Landa, Spanish Institute of Oceanography (IEO), Spain</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Josipa Ferri, <email xlink:href="mailto:jferri@unist.hr">jferri@unist.hr</email>; Marco Albano, <email xlink:href="mailto:malbano@unime.it">malbano@unime.it</email>
</p>
</fn>
<fn fn-type="equal" id="fn003">
<p>&#x2020;These authors have contributed equally to this work</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>01</day>
<month>05</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>11</volume>
<elocation-id>1347897</elocation-id>
<history>
<date date-type="received">
<day>01</day>
<month>12</month>
<year>2023</year>
</date>
<date date-type="accepted">
<day>02</day>
<month>04</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2024 D&#x2019;Iglio, Famulari, Ferri, Albano, Span&#xf2;, Capillo and Savoca</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>D&#x2019;Iglio, Famulari, Ferri, Albano, Span&#xf2;, Capillo and Savoca</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Improve the knowledge base on the ecomorphological adaptation of teleost species to different environments, trying to reconstruct how habitat can shape <italic>sagittae</italic>, is essential for conservational purposes, evolutionary evaluations, and population dynamics studies. Here is provided a comparative study between <italic>sagittae</italic> features, growth rates and age composition of two <italic>Scorpaena porcus</italic> populations inhabiting the Mediterranean waters, one from the Strait of Messina (Central Mediterranean Sea) and one from the Split area (Adriatic Sea). Additionally, the stomach contents of the Messina specimens have also been analysed to provide some preliminary information on the diet of <italic>S. porcus</italic>. A total of ninety individuals, half from each area, have been obtained from two extremely different environments. Results showed an overall different morphology, shape, and morphometry of <italic>sagittae</italic> among the size classes of the two investigated populations. Samples from Messina were characterized by a most elliptical and slender shape, with a more regular serration of margins than those from Split, which exhibited a wider <italic>sagitta</italic>, with a most enhanced <italic>anti-rostrum</italic> and longer <italic>rostrum</italic>. Concerning diet, specimens from Messina showed a preference for Crustacea (especially Brachyura and Amphipoda) and Teleost species, showing some differences with literature data from other geographical areas (Split one included). Results have confirmed the reliability of <italic>sagittae</italic> to detect the inter-population variability of <italic>S. porcus</italic> from different geographical areas, an essential tool for stock assessment, population studies and investigation on ecomorphological adaptation of teleost species to different habitats.</p>
</abstract>
<kwd-group>
<kwd>otolith</kwd>
<kwd>populations</kwd>
<kwd>growth</kwd>
<kwd>comparison</kwd>
<kwd>morphology</kwd>
<kwd>life-history</kwd>
</kwd-group>
<counts>
<fig-count count="11"/>
<table-count count="4"/>
<equation-count count="0"/>
<ref-count count="145"/>
<page-count count="20"/>
<word-count count="10585"/>
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<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Marine Biology</meta-value>
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</custom-meta-wrap>
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</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>The family Scorpaenidae counts more than 1400 species and represents one of the most diverse fish groups worldwide (<xref ref-type="bibr" rid="B128">Turan et&#xa0;al., 2009</xref>). It includes species adapted to live in many different environments, from the coastal shallow waters to the deep seas (<xref ref-type="bibr" rid="B29">Costello et&#xa0;al., 2001</xref>). Within the family, species belonging to <italic>Scorpaena</italic> genus are among the most abundant in all the temperate marine environments, and in several tropical seas, around the world, with 61 valid species included in the genus (<xref ref-type="bibr" rid="B96">Nelson et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B47">Fricke et&#xa0;al., 2018</xref>).</p>
<p>Among these, the black scorpionfish, <italic>Scorpaena porcus</italic> (Linnaeus, 1758), is one of the most common in the entire Mediterranean basin, Black Sea, and the Eastern Atlantic Ocean, from the British Islands to the Atlantic Moroccan coasts (<xref ref-type="bibr" rid="B141">Wheeler et&#xa0;al., 1986</xref>; <xref ref-type="bibr" rid="B88">Mah&#xe9; et&#xa0;al., 2014</xref>). It dwells rocky, or mixed rocky-sandy, benthic habitats, from the shallow water to the bathyal plan, with a bathymetric distribution in the Mediterranean basin ranging between 20 and 200 m (up to 800 m) with a mean at 20 m (<xref ref-type="bibr" rid="B47">Fricke et&#xa0;al., 2018</xref>). It is very common also on seagrass beds, showing a sedentary lifestyle with nocturnal feeding habits (<xref ref-type="bibr" rid="B112">Poutiers, 1987</xref>; <xref ref-type="bibr" rid="B103">Pashkov et&#xa0;al., 1999</xref>). According to literature, this species is a small-body benthic predator, with a preference for small fishes and crustaceans (<xref ref-type="bibr" rid="B26">Compaire et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B8">Aydin and Mazlum, 2020</xref>; <xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>). It is considered, together with the other scorpionfish species, as an essential predator for the well-functioning of the rocky-reefs ecosystems worldwide, both in temperate and tropical areas. Unlike other congeneric species, <italic>S. porcus</italic> is characterized by a slow-growth and a relatively short life, with 12 years as maximum recorded age (<xref ref-type="bibr" rid="B75">Kutsyn et&#xa0;al., 2019</xref>), and an enhanced geographical heterogeneity in life span and growth rates within the distribution areas (<xref ref-type="bibr" rid="B64">Jardas and Pallaoro, 1992</xref>; <xref ref-type="bibr" rid="B103">Pashkov et&#xa0;al., 1999</xref>; <xref ref-type="bibr" rid="B76">Kuzminova et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B44">Ferri et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B75">Kutsyn et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B115">Sahin et&#xa0;al., 2019</xref>). This high inter population variability has been related to the influence of the different fishing pressures and environmental factors experienced by the populations within the distribution range (<xref ref-type="bibr" rid="B17">Bradai and Bouain, 1988</xref>; <xref ref-type="bibr" rid="B64">Jardas and Pallaoro, 1992</xref>; <xref ref-type="bibr" rid="B135">Unsal and Oral, 1994</xref>; <xref ref-type="bibr" rid="B78">La Mesa et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B75">Kutsyn et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B115">Sahin et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>). Many authors have also suggested <italic>S. porcus</italic> as indicator species for biomonitoring and environmental conditions&#x2019; assessments (<xref ref-type="bibr" rid="B114">Rudneva et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B89">Medinets and Medinets, 2010</xref>; <xref ref-type="bibr" rid="B76">Kuzminova et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B44">Ferri et&#xa0;al., 2012</xref>). In the Black Sea, this species showed a significant negative trend in body size related to several factors, as pollution, and temperature rise, being negatively affected by anthropogenetic pressure (<xref ref-type="bibr" rid="B114">Rudneva et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B89">Medinets and Medinets, 2010</xref>; <xref ref-type="bibr" rid="B76">Kuzminova et&#xa0;al., 2011</xref>). While, in the Adriatic Sea, variations in population dynamics, biological traits and biomass, related to fishing pressure, confirmed the sensitivity of the species to fisheries activities (<xref ref-type="bibr" rid="B65">Jardas et&#xa0;al., 1998</xref>; <xref ref-type="bibr" rid="B63">Jardas, 1999</xref>; <xref ref-type="bibr" rid="B101">Oven et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B124">Stagli&#x10d;i&#x107; et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B44">Ferri et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>). <italic>S. porcus</italic> represents one of the main target species of Mediterranean artisanal fisheries (especially in the eastern and western parts of the basin), representing a large portion of the total catch and, consequently, a large source of income (<xref ref-type="bibr" rid="B63">Jardas, 1999</xref>; <xref ref-type="bibr" rid="B48">Garc&#xed;a-Rodr&#xed;guez et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B46">Forcada Almarcha et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B50">Go&#xf1;i et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B45">Forcada et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B78">La Mesa et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B44">Ferri et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B102">&#xd6;zg&#xfc;l et&#xa0;al., 2019</xref>) thanks to its relatively high commercial value in Croatia, Spain, and Italy. <italic>S. porcus</italic> annual total catch was 240 tons in the Eastern part (192 tons in Turkey) of the basin (104 tons from Black Sea, 36 from the Sea of Marmara, 88 from Aegean Sea and 12 from the Mediterranean Sea) in the first decade of 2000 (<xref ref-type="bibr" rid="B14">Bilgin and &#xc7;elik, 2009</xref>). In Italy the total landings of scorpionfish species in 2021 changed in relation to the geographic areas, from 41.2 tons in the Adriatic Sea, to 446.58 tons in the Ionia Sea (<xref ref-type="bibr" rid="B42">FAO, 2020</xref>). Concerning <italic>S. porcus</italic> conservation status, it was recorded a relevant decline in catches and average size during the last two decades in the Ionian coast of the Sicily, strictly related to fishing activities (<xref ref-type="bibr" rid="B127">Tiralongo, 2024</xref>). To improve the knowledge base on the intra-specific variability of <italic>S. porcus</italic>, analyzing the different growth and eco-morphological adaptations of populations inhabiting different geographical areas, is essential to enhance its conservation. In fact, in fishery biology, a proper assessment of fishes&#x2019; populations dynamics and stocks&#x2019; structure is a fundamental step to establish good management measures, monitoring species and communities&#x2019; response to management actions and different exploitation levels (<xref ref-type="bibr" rid="B24">Charles, 2001</xref>; <xref ref-type="bibr" rid="B25">Cochrane, 2002</xref>; <xref ref-type="bibr" rid="B39">Dimech et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B108">Perdichizzi et&#xa0;al., 2022</xref>). Moreover, being <italic>S. porcus</italic> a benthic species with a low home-range, a high spatial heterogeneity, and an enhanced ecological value, investigate how different habitats can influence its otoliths&#x2019; features (such as shape, morphology, and morphometry) and growth rates acquires great relevance and scientific interest, also in terms of taxonomical studies and evolution.</p>
<p>
<italic>Sagittae</italic>, as the other otoliths, grow for the entire fishes&#x2019; life, with a daily metabolically inert deposition of calcium carbonate (<xref ref-type="bibr" rid="B110">Popper et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B126">Thomas and Swearer, 2019</xref>). Their peculiar physiology and growing mechanism have made otoliths (mainly <italic>sagittae</italic>) (<xref ref-type="bibr" rid="B4">Assis, 2003</xref>, <xref ref-type="bibr" rid="B5">2005</xref>; <xref ref-type="bibr" rid="B77">Ladich and Schulz-Mirbach, 2016</xref>) an essential tool to study fish ages and growth (<xref ref-type="bibr" rid="B98">Newman and Dunk, 2002</xref>; <xref ref-type="bibr" rid="B136">Velasco et&#xa0;al., 2011</xref>). Thanks to their high variability in morphology and shape between and within species (<xref ref-type="bibr" rid="B140">Volpedo et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B62">Jaramilo et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B143">Zhuang et&#xa0;al., 2015a</xref>; <xref ref-type="bibr" rid="B34">D&#x2019;Iglio et&#xa0;al., 2021a</xref>; <xref ref-type="bibr" rid="B37">D&#x2019;iglio et&#xa0;al., 2022a</xref>; <xref ref-type="bibr" rid="B36">D&#x2019;Iglio et&#xa0;al., 2023</xref>), they have been widely used also in several research field (e.g., taxonomy, paleontological studies, stomach content analyses, and stocks assessment in fisheries studies) (<xref ref-type="bibr" rid="B12">Benzinou et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B55">Higgins et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B95">Muniz et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B97">Neves et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B35">D&#x2019;Iglio et&#xa0;al., 2021b</xref>; <xref ref-type="bibr" rid="B38">D&#x2019;Iglio et&#xa0;al., 2022b</xref>). Concerning the intra-specific variations in shape and morphology, they can be related to many factors, ranging from those genetically driven (<xref ref-type="bibr" rid="B138">Vignon and Morat, 2010</xref>), to environmental conditions and diet (<xref ref-type="bibr" rid="B120">Schulz-Mirbach et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B55">Higgins et&#xa0;al., 2013</xref>). For this reason, <italic>sagittae</italic> are considered a good phenotypic marker, reliable to investigate the ecomorphological adaptation of teleost species to different environmental conditions and ecological dynamics (<xref ref-type="bibr" rid="B1">Abaad et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B16">Bose et&#xa0;al., 2020</xref>), as highlighted by the shape and morphological differences between stocks and populations (<xref ref-type="bibr" rid="B12">Benzinou et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B55">Higgins et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B95">Muniz et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B97">Neves et&#xa0;al., 2021</xref>). Diet composition and feeding habits were assessed as factors which influence on teleosts physiology, metabolism, and, consequently, on otoliths too (<xref ref-type="bibr" rid="B93">Mommsen, 1998</xref>; <xref ref-type="bibr" rid="B116">Sanchez-Jerez et&#xa0;al., 2002</xref>). Moreover, molecules incoming deriving from diet strongly influence otoliths&#x2019; growth process and the geographical differences in feeding habits, added to genetic and environmental variability, could allow their inter-population variability (<xref ref-type="bibr" rid="B91">Mille et&#xa0;al., 2016a</xref>, <xref ref-type="bibr" rid="B92">2016b</xref>; <xref ref-type="bibr" rid="B58">H&#xfc;ssy et&#xa0;al., 2021</xref>).</p>
<p>The present paper aims to investigate the intra-specific differences in age composition, growth, diet composition, and <italic>sagittae</italic> shape and morphology, between two populations of <italic>S. porcus</italic> inhabiting, respectively, the tidal ponds in the beach rock formations, along the Strait of Messina coast (Central Mediterranean Sea), and the coastal environment near the Split Area (Eastern Adriatic Sea). The stomach content analysis has been performed only on the specimens from Messina, using the obtained data to make a comparison with the data present in literature for the diet composition of the studied species in the Split area (<xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>). The main purposes were: (i) to assess how <italic>sagittae</italic> contours, morphology and morphometry change among size classes in the two studied populations; (ii) to evaluate the reliability of shape and morphometrical analysis in the assessment of different <italic>S. porcus</italic> populations; (iii) to provide preliminary observations on growth and age composition between the studied areas; (iv) to analyse the stomach contents of the specimens from Messina, providing preliminary data on the diet composition and feeding habits of the samples of this area, useful to improve the knowledge base on their geographical variability and their influence on the growth and age composition of the specimens from the two different studied areas. These information are important to improve the knowledge base on the influences of the environment on inner ear morphology, otoliths and somatic growth.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Sample collection and studied areas.</title>
<p>A total number of 90 <italic>S. porcus</italic> specimens were collected in December 2022 from two different localities of the Mediterranean Sea: 45 specimens from the tidal ponds occurring toward the Sicilian coast of the Messina&#x2019;s Strait (Central Mediterranean Sea, 38&#xb0;15&#x2019;25&#x201d;N, 15&#xb0;36&#x2019;51&#x201d;E), and 45 specimens from the coastal environment near the Split area (Eastern Adriatic Sea, 43&#xb0;26&#x2019;0&#x201d;N, 16&#xb0;26&#x2019;55&#x201d;E) (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Map of the central Mediterranean Sea with the two studied areas highlighted in the boxes (Strait of Messina area in green and Split area in red).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g001.tif"/>
</fig>
<p>Thanks to the limited depth (ranging from 0.2 to 1.2 m) of the tidal ponds occurring in the &#x201c;beach rock&#x201d; formations, a coastal biotope extending in the north-eastern Messina coast, between the two villages &#x201c;Ganzirri&#x201d; and &#x201c;Torre Faro&#x201d;, toward the Sicilian coast of the Strait of Messina, specimens were caught manually, using a sampling net from the rocky bench present in the beachfront, by local fishermen.</p>
<p>&#x201c;Beach rock&#x201d;. This sedimentary formation represents a rocky bench (2 kilometres long) that arrive from the beachfront, in the intertidal zone, to a depth of 2-3 meters, at the beginning of the infralittoral zone. It is considered an area of high interest, being part of the Oriented Natural Reserve of &#x201c;Capo Peloro Lagoon&#x201d; (<xref ref-type="bibr" rid="B117">Savoca et&#xa0;al., 2020</xref>), which, thanks to its irregular shape, is characterized by several tidal ponds, with different surface and connection dynamics to the sea, hosting very complex ecological communities. These are essential nursery areas for several teleost species and a shelter zone for numerous taxa (<xref ref-type="bibr" rid="B52">Gravem and Morgan, 2017</xref>; <xref ref-type="bibr" rid="B19">Capillo et&#xa0;al., 2018</xref>).</p>
<p>Concerning specimens from the Split area (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>), they were collected using sets of trammel nets, 1.5 m high and 32 m long, with an inner layer mesh size of 28 mm and 150 mm mesh size of outer layers. Fish samples were captured in the nearshore, coastal waters in the Split area (43.5&#xb0;N), in the eastern Adriatic, at depths ranging between 10 and 40 m (20 m in average). This biotope is characterized mainly by rocky substrata covered by photophilic algae alternating with patches of sand and <italic>Posidonia oceanica</italic> seagrass beds (<xref ref-type="bibr" rid="B44">Ferri et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>).</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Samples processing, otolith extraction and age reading</title>
<p>All sampled specimens were transported frozen to the laboratory, where they were measured (TL, cm) and weighed (BW, g). <italic>Sagittae</italic> were collected from each specimen, cleaned (15 min in 3% H<sub>2</sub>O<sub>2</sub>, followed by Milli-Q water) and, once dried, stored in Eppendorf microtubes. To assess the intra-specific variability in <italic>sagittae</italic> shape and morphometries in relation to fish size, all samples were divided into three size classes according to the following TL values: Class I included specimens smaller than 100 mm, Class II those with a TL ranging from 100 mm to 180 mm, and Class III larger than 180 mm. The left <italic>sagittae</italic> of both populations were photographed on the distal side using a stereomicroscope (Olympus SZX10) equipped with an Olympus DP-25 digital camera. Images were converted in binary format for shape analysis, using the ImageJ 1.48p free software, &#x201c;available at <ext-link ext-link-type="uri" xlink:href="http://rsb.info.nih.gov/ij/">http://rsb.info.nih.gov/ij/</ext-link>&#x201d;. Otoliths morphologies were described following the terminology used by Tuset (<xref ref-type="bibr" rid="B131">Tuset et&#xa0;al., 2008</xref>), Nolf (<xref ref-type="bibr" rid="B99">Nolf, 1985</xref>) and Assis (<xref ref-type="bibr" rid="B3">Assis, 2000</xref>).</p>
<p>After images collection, left <italic>sagittae</italic> were used for age estimation. The reading process was carried out by two different operators, without data on fish length, one with experience in estimating the age of <italic>S. porcus</italic>, and one with experience in estimating the age of other teleost species. Both readers performed two different readings, with one month of distance one from the other. If readings were different for one or two years, it was performed a third reading, with the otolith discarding in the case of persisting differences. The relative precision between readings was tested by the index of average percentage error (APE) (<xref ref-type="bibr" rid="B11">Beamish and McFarlane, 1983</xref>) and the coefficient of variation (CV) (<xref ref-type="bibr" rid="B22">Chang, 1982</xref>). The annual age of the specimens was estimated with reflected light and on a black background using the stereomicroscope (Olympus SZX10) equipped with an Olympus DP-25 digital camera, at magnification 1.5-2.4x). The pattern of opaque and translucent zones was used for annuli counting. Indeed, according to literature (<xref ref-type="bibr" rid="B78">La Mesa et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B20">Carbonara and Follesa, 2019</xref>; <xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>), an annulus is the combination of an opaque and a translucent (or hyaline) ring, appearing, under the reflecting light respectively light and dark. When this pattern was not clear, <italic>sagittae</italic> were grinded to facilitate the rings&#x2019; counting. It was not necessary the otoliths sectioning, being grinding enough to reveal the opaque and translucent zones&#x2019; pattern for those samples in which this was not clear from the whole otoliths viewing. Age was estimated by counting translucent zones (<xref ref-type="bibr" rid="B79">Landa and Hern&#xe1;ndez, 2020</xref>) and considering the 1<sup>st</sup> of July as the birthdate (<xref ref-type="bibr" rid="B78">La Mesa et&#xa0;al., 2010</xref>). In addition, during the age reading process, the edge of otoliths has been classified as opaque or translucent.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Stomach content analysis</title>
<p>Concerning stomachs, they were sampled only from Messina&#x2019;s specimens, and stored in ethanol 70% + glycerin 5% for stomach content analysis. Each stomach content of specimens from Messina was analyzed under the stereomicroscope, to identify, at the lowest taxonomic level possible, each prey. Each preys&#x2019; items were counted and weighted, also detecting the digestion degree (1 = undamaged; 2 = almost digested; 3 = highly digested). The anatomical undigested preys&#x2019; parts (e.g., otoliths, telsons, carapaces, mouth parts, heads capsules, fish columns) were counted to assess the contribution of each prey taxon to the diet, grouping the unidentifiable preys&#x2019; items, due to the advanced state of digestion, into undetermined taxa. According to their presence in the stomachs, only one type of anatomical remains was counted for each prey group to avoid the double-counting.</p>
<p>Following indexes were calculated to evaluate the contribution of each preys&#x2019; taxa to the diet: the percentage of biomass composition (%W), the percentage of abundance composition (%<italic>N</italic>), and the frequency of occurrence (%F) (<xref ref-type="bibr" rid="B59">Hyslop, 1980</xref>). The Relative Importance Index was also calculated [IRI = %F (%N + %P)] expressing it also as percentage 9<inline-formula>
<mml:math display="inline" id="im1">
<mml:mrow>
<mml:mtext>%&#xa0;IRI</mml:mtext>
<mml:mo>=</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mrow>
<mml:mtext>IRI</mml:mtext>
</mml:mrow>
<mml:mtext>i</mml:mtext>
</mml:msub>
<mml:msubsup>
<mml:mrow>
<mml:mtext>&#xa0;&#x3a3;</mml:mtext>
</mml:mrow>
<mml:mi>i</mml:mi>
<mml:mtext>N</mml:mtext>
</mml:msubsup>
<mml:msub>
<mml:mrow>
<mml:mtext>&#xa0;IRI</mml:mtext>
</mml:mrow>
<mml:mtext>i</mml:mtext>
</mml:msub>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mtext>&#xa0;</mml:mtext>
<mml:mo>&#xd7;</mml:mo>
<mml:mtext>&#xa0;</mml:mtext>
<mml:mn>100</mml:mn>
</mml:mrow>
</mml:math>
</inline-formula>)(<xref ref-type="bibr" rid="B27">Cort&#xe9;s, 1997</xref>; <xref ref-type="bibr" rid="B35">D&#x2019;Iglio et&#xa0;al., 2021b</xref>; <xref ref-type="bibr" rid="B38">D'Iglio et al., 2022b</xref>). Concerning the empty stomachs, it was calculated the Vacuity Index, VC = (Ne/N) &#xd7; 100, as the percentage of empty stomachs (Ne) on the total stomach number (N).</p>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Shape and morphometric analysis</title>
<p>Shape R (R software package, RStudio 2022.07.1 Build 554; R Gui 4.1.3 2022.03.10) was used to perform the Shape analysis from the otoliths&#x2019; outlines. This package has been developed for inter and inter specific analysis on shape variability in teleost&#x2019;s otoliths (<xref ref-type="bibr" rid="B81">Libungan and P&#xe1;lsson, 2015</xref>). Each binarized <italic>sagitta</italic> photo was analyzed for the outlines&#x2019; detection using a shape R specific function, with 0.05 as value for intensity threshold greyscale. A data file with studied specimens&#x2019; information (as body weight and fish length) was linked to extracted contours. These was used for each size classes to perform several otoliths&#x2019; measurements (maximum width, OW, mm, maximum length, OL, mm, perimeter, OP, mm, and surface, OS, mm<sup>2</sup>), through the getMeasurements function. Proper package functions were applied for the extraction, and subsequent adjusting, of Wavelet and Fourier coefficients, to assess the allometric relationships between fish lengths and otolith shapes. Wavelet coefficients were used to provide the comparison between the mean <italic>sagittae</italic> shape of the analyzed populations. The reconstruction&#x2019;s quality was estimated analyzing the deviation of the coefficient&#x2019;s reconstruction from the otolith outline (<xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Figure S1</bold>
</xref>). Finally, a g-plots R package&#x2019;s specific function was used to investigate how the position along the outline can influence the wavelet coefficients variation (<xref ref-type="supplementary-material" rid="SF2">
<bold>Supplementary Figure S2</bold>
</xref>).</p>
<p>Otoliths measurements performed with shape R were used to calculate several indexes. The <italic>sagittae</italic> length increase, related to the total fish length, was evaluated assessing otolith length to total fish length ratio (OL/TL); while, according to literature (<xref ref-type="bibr" rid="B132">Tuset et&#xa0;al., 2003a</xref>, <xref ref-type="bibr" rid="B133">2003b</xref>, <xref ref-type="bibr" rid="B129">2016a</xref>; <xref ref-type="bibr" rid="B105">Pavlov, 2016</xref>, <xref ref-type="bibr" rid="B106">2021</xref>; <xref ref-type="bibr" rid="B66">Jawad et&#xa0;al., 2018</xref>), several shape indexes were used to evaluate intra and inter specific variability of <italic>sagittae</italic> shape: roundness (Ro = 4OS/&#x3c0;OL<sup>2</sup>), form factor (FF = 4&#x3c0;OS/OP<sup>2</sup>), aspect ratio (AR = OW/OL%), circularity (C = OP<sup>2</sup>/OS), rectangularity (Re = OS/[OL&#xd7;OW]) and ellipticity [E = (OL&#x2013;OW)/(OL+OW)].</p>
</sec>
<sec id="s2_5">
<label>2.5</label>
<title>Statistical analysis</title>
<p>Length frequency distributions of individuals from the two sampling areas were compared using the Kolmogorov&#x2013;Smirnov two-sample test.</p>
<p>The von Bertalanffy growth model was fitted to the estimated age-length dataset using a non-linear least-square procedure of a Gauss&#x2013;Newton algorithm, included in the statistical package STATISTICA (version 14.0.0.15). The von Bertalanffy growth parameters (L<sub>&#x221e;</sub>, K and t<sub>0</sub>) were calculated for both fish populations sampled in the Messina&#x2019;s Strait and Split area and compared by the multivariate Hotelling&#x2019;s T<sup>2</sup>-test. The growth performance index (&#x424;&#x2019; = 2 log L&#x221e; + log K) was then calculated to compare the different populations of the black scorpionfish throughout its distribution range.</p>
<p>Univariate and multivariate statistical methods were applied to conduct investigations on <italic>sagittae</italic> features of specimens from Messina and Split using Prism V.8.2.1 (Graph- pad Software Ltd., La Jolla, CA 92037, USA), R vegan package V.2.5, and PAST V.4.</p>
<p>The intra-population variability of <italic>sagittae</italic> morphometries, between the different size classes investigated, was detected using a one-way analysis of variance (one-way ANOVA) or Kruskal&#x2013;Wallis one-way ANOVA, followed by Tukey or Dunn&#x2019;s post-hot test respectively. A Principal Component Analysis (PCA) was conducted to obtain an overview of the differences in otolith parameters between the size classes examined within each population. Additionally, the correlation between the measured parameters and fish body weight (BW) and total length (TL) was tested using the Spearman correlation analysis. To explore the intra-specific variability of otolith contours within each population, the shape indices were extrapolated and analysed through an ANOVA-like permutation test and a PCA, to obtain an overview of the differences in otolith shape between the size classes examined at the intra-population level. The significance level of p-value was set at &lt; 0.05.</p>
<p>Concerning the inter-population variability, a one-way analysis of variance, or a corresponding non-parametric test in case of non-homogeneity of the data, was used to verify the existence of any differences in the <italic>sagittae</italic> extracted between Messina and Split specimens, using the origin site of the samples as the only independent factor, followed by a PCA. For a more detailed description of the two populations examined, a Mann-Whitney test was used to explore the variability of individual otolith morphometric parameters, within each size class, between the sampling sites identified in the current study. Additionally, to obtain an overview of the <italic>sagittae</italic> features of the two populations under examination, a Linear Discriminant Analysis (LDA) was applied to the dataset. A cross-validation of the LDA was carried out to provide a classification of the individuals for both the investigated areas, assessing the classification success rate as the specimens&#x2019; percentage and number correctly assigned to the original sample. Finally, to investigate the inter-population variability of otolith contours between specimens from Messina and Split, the shape indices were extrapolated and analysed through an ANOVA-like permutation test and a Principal Component Analysis (PCA), to obtain an overview of the differences in otolith shape between the two analyzed populations.</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<p>The 45 specimens from Messina showed a TL ranging from 6.5 to 23 cm, with a Mean TL &#xb1; SD of 12.61 &#xb1; 0.69 cm. Concerning Split area, specimens showed a TL ranging from 11.2 to 24.1 cm and a Mean TL &#xb1; SD of 14.71 &#xb1; 0.41 cm. Length-frequency distributions were significantly different between the two sampling areas (Kolmogorov&#x2013;Smirnov test; P &lt; 0.001).</p>
<sec id="s3_1">
<label>3.1</label>
<title>Growth rates and age composition</title>
<p>Age counts were successfully provided for all the otoliths from Messina and Split, with no sample discarded, and a quite low variability of APE and CV indices (4.7% and 9.4%, respectively). The inner structure of sagittal otoliths consisted of a wide opaque nucleus, surrounded by an alternating pattern of translucent and opaque zones. Interpretation of otolith edge types revealed that translucent edge type was dominant in samples from Messina and Split (&gt;90% in both areas). The age composition by area (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>) showed 3 to 5 years old as the main estimated ages (57.77%) for the Messina specimens, with a maximum age of 5 years. In Split area, the larger part of the specimens was between 5 and 8 years old (75.57%), with a maximum age of 8 years, showing that the specimens inhabiting the Messina&#x2019;s tidal ponds was younger than that from Split.</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Age-length keys and Lenght-at-Age of <italic>S. porcus</italic> specimens from Messina and Split area.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="center"/>
<th valign="top" colspan="5" align="center">Messina</th>
<th valign="top" colspan="7" align="center">Split</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="center">TL (cm)</td>
<td valign="top" align="center">1</td>
<td valign="top" align="center">2</td>
<td valign="top" align="center">3</td>
<td valign="top" align="center">4</td>
<td valign="top" align="center">5</td>
<td valign="top" align="center">2</td>
<td valign="top" align="center">3</td>
<td valign="top" align="center">4</td>
<td valign="top" align="center">5</td>
<td valign="top" align="center">6</td>
<td valign="top" align="center">7</td>
<td valign="top" align="center">8</td>
</tr>
<tr>
<td valign="top" align="center">7</td>
<td valign="top" align="center">7</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">8</td>
<td valign="top" align="center">6</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">9</td>
<td valign="top" align="center">5</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">10</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">11</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">2</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">12</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center">4</td>
<td valign="top" align="center">3</td>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">13</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">2</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">5</td>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">14</td>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center">6</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center">3</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">15</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">2</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
<td valign="top" align="center">7</td>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">16</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">3</td>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">3</td>
<td valign="top" align="center">3</td>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">17</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">5</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">2</td>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">18</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">19</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center">2</td>
</tr>
<tr>
<td valign="top" align="center">20</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">21</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
</tr>
<tr>
<td valign="top" align="center">22</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">23</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">2</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">24</td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center">1</td>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="center">N</td>
<td valign="top" align="center">18</td>
<td valign="top" align="center">1</td>
<td valign="top" align="center">22</td>
<td valign="top" align="center">2</td>
<td valign="top" align="center">2</td>
<td valign="top" align="center">1</td>
<td valign="top" align="center">4</td>
<td valign="top" align="center">6</td>
<td valign="top" align="center">7</td>
<td valign="top" align="center">17</td>
<td valign="top" align="center">7</td>
<td valign="top" align="center">3</td>
</tr>
<tr>
<td valign="top" align="center">%</td>
<td valign="top" align="center">40</td>
<td valign="top" align="center">2.22</td>
<td valign="top" align="center">48.89</td>
<td valign="top" align="center">4.44</td>
<td valign="top" align="center">4.44</td>
<td valign="top" align="center">2.22</td>
<td valign="top" align="center">8.89</td>
<td valign="top" align="center">13.33</td>
<td valign="top" align="center">15.56</td>
<td valign="top" align="center">37.78</td>
<td valign="top" align="center">15.56</td>
<td valign="top" align="center">6.67</td>
</tr>
<tr>
<td valign="top" align="center">Length-at-Age (cm)</td>
<td valign="middle" align="center">7.77</td>
<td valign="middle" align="center">14.00</td>
<td valign="middle" align="center">15.34</td>
<td valign="middle" align="center">17.25</td>
<td valign="middle" align="center">20.75</td>
<td valign="middle" align="center">11.90</td>
<td valign="middle" align="center">11.77</td>
<td valign="middle" align="center">12.30</td>
<td valign="middle" align="center">13.67</td>
<td valign="middle" align="center">14.82</td>
<td valign="middle" align="center">17.54</td>
<td valign="middle" align="center">19.53</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>The von Bertalanffy growth curves were fitted to age-length dataset, estimated for individuals from the Messina Strait and Split area (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>). Estimated parameters for <italic>S. porcus</italic> from the Messina Strait and Split area were L<sub>&#x221e;</sub> = 28.78 cm, K = 0.22 year<sup>-1</sup>, t<sub>0</sub> = -0.02 year (R<sup>2 </sup>= 0.89) and L<sub>&#x221e;</sub> = 40.59 cm, K = 0.05 year<sup>-1</sup>, t<sub>0</sub> = -2.58 year (R<sup>2 </sup>= 0.73), respectively. The growth performance index (&#x424;&#x2019;) was 2.26 for the fish from the Messina Strait and 1.92 for the fish from the Split area. Specimens from Split Area showed a growth rate of slightly more than 1 cm per year between 4 and 7 years old.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Fitted von Bertalanffy growth curve for S. porcus individual from Messina and Split.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g002.tif"/>
</fig>
<p>The Hotelling&#x2019;s T<sup>2</sup>-test indicated that the von Bertalanffy growth curves differed significantly between the two tested groups of individuals (T<sup>2 </sup>= 2885.92 &gt; T<sup>2</sup>
<sub>0 (0.05, 3,86)</sub> = 12.20). The L<sub>&#x221e;</sub> value was higher in fish from the Split area and the K value was higher in fish from the Messina Strait. Therefore, considering the growth coefficient and the growth performance index, fish from the Messina Strait grew faster than fish from the Split area. Statistical difference in length-at-age data (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>) was observed between the two investigated areas (t-test for paired comparison, P = 0.025).</p>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Morphometric and shape analysis</title>
<p>Studied specimens showed an overall morphology of <italic>sagittae</italic> characterized by an oblong to lanceolate outline, with serrate to crenate margins. The anterior region was peaked, with <italic>rostrum</italic>, <italic>antirostrum</italic> and <italic>excisura ostii</italic>, that, together with the notch tilt angle, changed between the two populations in the analyzed size classes (<xref ref-type="fig" rid="f3">
<bold>Figures&#xa0;3A&#x2013;E</bold>
</xref>). The posterior region was oblique to irregular, and the <italic>sulcus acusticus</italic> was heterosulcoid, ostial and median.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Stereoscope images of the medial view of the sagittae belonging to specimens from Messina <bold>(A-C)</bold> and Split <bold>(D, E)</bold> for the three investigated size classes [Class I: <bold>(A)</bold> Class II: <bold>(B&#x2013;D)</bold> Class III: <bold>(C&#x2013;E)</bold>]. Red dots indicate the end of each hyaline annulus in the reading area of each otoliths&#x2019; image.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g003.tif"/>
</fig>
<p>In <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Tables S1</bold>
</xref>, <xref ref-type="supplementary-material" rid="SM1">
<bold>S2</bold>
</xref> are provided the morphometric mean values of the left <italic>sagittae</italic> for the two investigated populations, with the minimum and maximum range, divided in the three size classes.</p>
<p>
<italic>Sagittae</italic> belonging to Messina specimens showed an oblong shape, with an enhanced margins serration. The <italic>rostrum</italic> and the <italic>antirostrum</italic> were short, with an arrow <italic>excisura ostii</italic>, a not acute notch, and an irregular posterior margin (<xref ref-type="fig" rid="f4">
<bold>Figures&#xa0;4A&#x2013;C</bold>
</xref>). Concerning the intra specific differences among size classes, morphometrical analyses showed an enhanced variability, confirmed also by the mean otoliths&#x2019; shapes (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4D</bold>
</xref>). <italic>Sagittae</italic> of specimens belonging to Size Class I showed a most circular contour, then the other classes. Also, the posterior region showed a peculiar organization, with a most oblique margin than the other size classes. In the Size Class II, the contour was less circular than the Class I, with most irregular posterior margin and less acute notch of <italic>excisura</italic>. The Size Class III showed a very different shape, with an enhanced ellipticity (as highlighted by the highest E value among size classes: 0.41 &#xb1; 0.02 mm, see <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S1</bold>
</xref>) resulting in a less wide and longer <italic>sagitta</italic> than the other size classes.</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Stereoscope images of the medial view of the sagittae belonging to specimens from Messina, for the three investigated size classes [<bold>(A)</bold> Class I, <bold>(B)</bold>&#xa0;Class II, <bold>(C)</bold> Class III], with their mean otoliths shape <bold>(D)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g004.tif"/>
</fig>
<p>Overall, the specimens examined showed significant variations in otoliths&#x2019; parameters between the size classes considered in the study.</p>
<p>The results of the differences are reported in <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S3</bold>
</xref>, showing the absence of significant differences for 90% of the parameters between Class II and Class III. Concerning the comparison between Size Class I and II, and between Size Class I and III, significant differences were detected for all the investigated morphometrical parameters, except for Re values.</p>
<p>PCA confirmed the results obtained by univariate analysis (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>), explaining the main differences observed between the classes I vs II and I vs III.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Principal Component Analysis (PCA) plot between the specimens belonging to size classes I (fill triangle dark magenta), II (dot orange red) and III (fill square forest green), calculated on sagittae parameters of Messina population <bold>(A)</bold>; Principal Component Analysis (PCA) plot between the Messina&#x2019;s specimens belonging to size classes I (fill triangle dark magenta), II (dot orange red) and III (fill square forest green), calculated on elliptic Fourier descriptors <bold>(B)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g005.tif"/>
</fig>
<p>All <italic>sagittae</italic> measurements showed a significant correlation with specimens&#x2019; parameters TL and BW. Results of Spearman correlation analyses are reported in <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S4</bold>
</xref>.</p>
<p>The mean shape of otoliths differed significantly between the three size classes investigated (p&lt; 0.001). Marked differences in otolith shape have also been confirmed by PCA, mainly between the classes I and III (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5B</bold>
</xref>).</p>
<p>Concerning <italic>sagittae</italic> belonging to Split specimens, they showed a lanceolate contour, with a marked irregularity of margins. The <italic>rostrum</italic> and <italic>antirostrum</italic> were long, with a wide <italic>excisura</italic> and a very acute notch (<xref ref-type="fig" rid="f6">
<bold>Figures&#xa0;6A, B</bold>
</xref>). Concerning the intra-specific differences among size classes, shape analysis showed a general uniform shape (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6C</bold>
</xref>). Class II was characterized by a more enhanced circular contour than the other class, with an oblique posterior margin and an acute notch. In Class III the <italic>sagittae</italic> were more elliptic than the Class II, as highlighted by the highest E value (0.41 &#xb1; 0.02 mm, see <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S2</bold>
</xref>), with an enhanced length and a reduced width.</p>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>Stereoscope images of the medial view of the sagittae belonging to specimens from Split, for the two investigated size classes [<bold>(A)</bold> Class II, <bold>(B)</bold> Class III], with their mean otoliths shape <bold>(C)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g006.tif"/>
</fig>
<p>The specimens examined showed overall a significant variation in <italic>sagittae</italic> parameters between the two size classes investigated in the study (p&lt;0.05).</p>
<p>The results of the variations observed for the individual parameters of the otoliths extracted from each size class are reported in <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S5</bold>
</xref>. Significant differences between the two size classes (Size Classes II and III) of the Split population were detected for all the investigated morphometrical parameters, except for Re values.</p>
<p>PCA confirmed the results obtained by univariate analysis (<xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7A</bold>
</xref>), explaining the main differences observed between the size.</p>
<fig id="f7" position="float">
<label>Figure&#xa0;7</label>
<caption>
<p>
<bold>(A)</bold> Principal Component Analysis (PCA) plot between the specimens belonging to size classes II (Blue fill inverted triangle) and III (Red fill diamond), calculated on sagittae parameters from Split population; <bold>(B)</bold> Principal Component Analysis (PCA) plot between the specimens belonging to size classes II and III, from Split calculated on elliptic Fourier descriptors.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g007.tif"/>
</fig>
<p>Almost all <italic>sagittae</italic> measurements showed significant correlation with specimen parameters TL and BW. Results of Spearman correlation analyses are reported in <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S6</bold>
</xref>.</p>
<p>The mean shape of otoliths differed significantly between the two size classes investigated (p&lt; 0.009). Slight differences in otolith shape have also been confirmed by PCA (<xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7B</bold>
</xref>).</p>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Geographical differences between sagittae features</title>
<p>Concerning the number of individuals compared between the two zones, specimens belonging to Class III were much lesser than those belonging to Class II in both the studied areas. Overall, despite quite an overlap between the two areas, the two groups of individuals examined were different both for fish parameters and <italic>sagittae</italic> features (P&lt;0.05), as well confirmed by PCA analysis (PC1 96.10%; PC2 3.51%) (<xref ref-type="fig" rid="f8">
<bold>Figure&#xa0;8</bold>
</xref>).</p>
<fig id="f8" position="float">
<label>Figure&#xa0;8</label>
<caption>
<p>Principal Component Analysis showing fish and sagittae features differences between the sampling sites: Messina (red dots) and Split (blue dots).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g008.tif"/>
</fig>
<p>Class I of the specimens from Messina was eliminated from subsequent analyses, as it was not comparable to a class equally represented in the other group from Split.</p>
<p>The results obtained through the Mann-Whitney test are reported in <xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>, showing, for the compared size Classes III, the presence of significant differences only for the Ro shape index.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Mann Whitney test results obtained for otoliths&#x2019; morphometric parameters comparison the same size classes from different sampling sites, Strait of Messina, and Split.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="bottom" align="center" rowspan="2">Sagittae parameters</th>
<th valign="middle" align="center">Messina <italic>vs</italic> Split Class II</th>
<th valign="middle" align="center">Messina <italic>vs</italic> Split Class III</th>
</tr>
<tr>
<th valign="bottom" align="center">P value</th>
<th valign="bottom" align="center">P value</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="bottom" align="center">
<bold>OS</bold>
</td>
<td valign="bottom" align="center">0.0116<sup>*</sup>
</td>
<td valign="bottom" align="center">0.3290 <sup>ns</sup>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>OL</bold>
</td>
<td valign="bottom" align="center">0.5636 <sup>ns</sup>
</td>
<td valign="bottom" align="center">0.2468 <sup>ns</sup>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>OW</bold>
</td>
<td valign="bottom" align="center">&lt;0.0001<sup>***</sup>
</td>
<td valign="bottom" align="center">0.1255 <sup>ns</sup>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>OP</bold>
</td>
<td valign="bottom" align="center">0.0328<sup>*</sup>
</td>
<td valign="bottom" align="center">0.4286 <sup>ns</sup>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>Ro</bold>
</td>
<td valign="bottom" align="center">&lt;0.0001<sup>***</sup>
</td>
<td valign="bottom" align="center">0.0043<sup>**</sup>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>FF</bold>
</td>
<td valign="bottom" align="center">0.5459 <sup>ns</sup>
</td>
<td valign="bottom" align="center">0.9307 <sup>ns</sup>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>E</bold>
</td>
<td valign="bottom" align="center">&lt;0.0001<sup>***</sup>
</td>
<td valign="bottom" align="center">0.5368 <sup>ns</sup>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>P<sup>2</sup>/A</bold>
</td>
<td valign="bottom" align="center">0.5459 <sup>ns</sup>
</td>
<td valign="bottom" align="center">0.9307 <sup>ns</sup>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>Re</bold>
</td>
<td valign="bottom" align="center">0.0382<sup>*</sup>
</td>
<td valign="bottom" align="center">0.7922 <sup>ns</sup>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>AR</bold>
</td>
<td valign="bottom" align="center">&lt;0.0001<sup>***</sup>
</td>
<td valign="bottom" align="center">0.5368 <sup>ns</sup>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>OL/TL</bold>
</td>
<td valign="bottom" align="center">0.0440<sup>*</sup>
</td>
<td valign="bottom" align="center">0.1255 <sup>ns</sup>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>P value legend: &gt;0.05, not significant (ns); 0.03*, moderately significant; 0.002 **, significant; 0.001 *** highly significant.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>As highlighted by the LDA (<xref ref-type="fig" rid="f9">
<bold>Figure&#xa0;9</bold>
</xref>), the major differences concerned size classes II. An overall successful classification rate of 83.12% was achieved, concerning the LDA, for both the size classes combined (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S7</bold>
</xref>). Specimens belonging to Class II showed the highest classifications, with values of 90.9% (from Messina) and 89.7% (from Split). Specimens belonging to Class III showed slightly lower values: 83.3% from Split and 80% from Messina.</p>
<fig id="f9" position="float">
<label>Figure&#xa0;9</label>
<caption>
<p>Linear Discriminant Analysis (LDA) between morphometric parameters (OL, OW, OP, OS, OL/TL, C, Re, E, AR, FF, Ro) of the investigated populations for the size classes II (orange-red dot Messina, Blue fill inverted triangle Split) and III (Forest green fill square Messina; Red diamond Split).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g009.tif"/>
</fig>
<p>The analysis of the <italic>sagittae</italic> contours of the two populations examined showed an evident variability between the specimens of class II (p=0.001) and class III (p=0.017). Results of PCA and mean otoliths shapes confirmed the differences observed for each class between the sampling sites, showing a variance of 41.75% (PC1) and 17.16% (PC2) for Class II, and 53.67% (PC1) and 17.5% (PC2) for Class III (<xref ref-type="fig" rid="f10">
<bold>Figures 10A, B</bold>
</xref>).</p>
<fig id="f10" position="float">
<label>Figure&#xa0;10</label>
<caption>
<p>Results of PCA performed on Wavelet coefficient obtained by sagittae contours of specimens collected from Messina (red dots) and Split (blue dots). Results are reported for Class II and III, figures <bold>(A, B)</bold> respectively. Wavelet coefficient variance for each size classes investigated is expressed as percentage. Comparison of the mean otoliths shapes from the two populations is provided in figures <bold>(C, D)</bold> respectively for Class II and III.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g010.tif"/>
</fig>
<p>In <xref ref-type="fig" rid="f10">
<bold>Figures&#xa0;10C, D</bold>
</xref> is provided a comparison of the mean otoliths&#x2019; contours for the three investigated size classes. Specimens from Class II showed visible inter-population differences regarding <italic>rostrum</italic> and overall mean contour, oval in individuals from Split and elliptical in those from Messina. In the Class III, the differences detected for the second size class were confirmed, with more marked <italic>excisura ostii</italic> showed by Split specimens than Messina ones, which reported a less marked irregularity of the margins, a shorter <italic>rostrum</italic> and a less pointed <italic>antirostrum</italic>.</p>
<p>The shape analysis performed on the total interclass samples showed mean <italic>sagittae</italic> shapes clearly different between the two analyzed populations (<xref ref-type="fig" rid="f11">
<bold>Figure&#xa0;11</bold>
</xref>). The main differences were detected in the <italic>rostrum</italic> organization (longer and more pointed in Split population than in the Messina one), in the <italic>excisura ostii</italic> (deeper in Split population than in the Messina one), in the <italic>antirostrum</italic> organization (more prominent and pointed in Split population than in the Messina one), and in the overall <italic>sagittae</italic> shape, oval in Split specimens and elliptical in Messina ones.</p>
<fig id="f11" position="float">
<label>Figure&#xa0;11</label>
<caption>
<p>Mean shape of sagittae contours of the investigated <italic>S. porcus</italic> population.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1347897-g011.tif"/>
</fig>
</sec>
<sec id="s3_4">
<label>3.4</label>
<title>Stomach content analysis</title>
<p>The analysis of the 49 sampled stomachs from Messina (9 empty stomachs, with a Vacuity Index of 4.9) has shown a total of 80 preys, belonging to 29 taxa. As reported in <xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref>, Crustacea and Osteichthyes were the major taxa with the highest number of preys, with also the highest values of IRI %. The infraorder Brachyura was the taxon which showed the highest IRI % value (IRI % = 32.81), followed by the species, belonging to the infraorder Brachuyra, <italic>Pachygrapsus marmoratus</italic> (Fabricius, 1787) (IRI % = 27.59), that was the taxon with the highest relative abundance (N % = 18.75) among the preys. The order Amphipoda (IRI % = 10.74) and the species <italic>Xantho pilipes</italic> (A. Milne-Edwards, 1867) (IRI % = 9.86), belonging to the infraorder Brachuyra, were the other two most relevant taxa for relative importance. Concerning the Osteichthyes preys, those belonging to the species <italic>Chelon auratus</italic> (Risso, 1810) (IRI % = 3.24) and <italic>Thalassoma pavo</italic> (Linnaeus, 1758) (IRI % = 2.36) were the most relevant preys, with the genus <italic>Parablennius</italic> as taxon with the highest relative abundance value (N % = 2.50). Concerning the class Polychaeta, it was the major taxa with the less relative importance (IRI % = 0.15).</p>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>Diet composition of <italic>S. porcus</italic> individuals with %N (relative abundance), %W (percentage in biomass), %F (frequency of occurrence), IRI (index of relative importance) and %IRI (index of relative importance expressed as percentage) expressed for each prey items.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">TAXA</th>
<th valign="middle" align="center">N%</th>
<th valign="middle" align="center">W%</th>
<th valign="middle" align="center">F%</th>
<th valign="middle" align="center">IRI</th>
<th valign="middle" align="center">IRI%</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">Polychaeta</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.04</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.044</td>
<td valign="top" align="center">0.15</td>
</tr>
<tr>
<td valign="middle" align="left">
<bold>Total Polychaeta</bold>
</td>
<td valign="top" align="center">
<bold>1.25</bold>
</td>
<td valign="top" align="center">
<bold>0.04</bold>
</td>
<td valign="top" align="center">
<bold>1.59</bold>
</td>
<td valign="top" align="center">
<bold>2.044</bold>
</td>
<td valign="top" align="center">
<bold>0.15</bold>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Xantho pilipes</italic>
</td>
<td valign="top" align="center">6.25</td>
<td valign="top" align="center">14.39</td>
<td valign="top" align="center">6.35</td>
<td valign="top" align="center">131.036</td>
<td valign="top" align="center">9.86</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Brachynotus sexdentatus</italic>
</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">1.74</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">4.742</td>
<td valign="top" align="center">0.36</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Pachygrapsus marmoratus</italic>
</td>
<td valign="top" align="center">18.75</td>
<td valign="top" align="center">14.24</td>
<td valign="top" align="center">11.11</td>
<td valign="top" align="center">366.524</td>
<td valign="top" align="center">27.59</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Pachygrapsus</italic> sp.</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">2.11</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">5.341</td>
<td valign="top" align="center">0.40</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Inachus dorsettensis</italic>
</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.30</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.464</td>
<td valign="top" align="center">0.19</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Percnon gibbesi</italic>
</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">1.74</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">4.742</td>
<td valign="top" align="center">0.36</td>
</tr>
<tr>
<td valign="middle" align="left">Brachyura</td>
<td valign="top" align="center">16.25</td>
<td valign="top" align="center">4.87</td>
<td valign="top" align="center">20.63</td>
<td valign="top" align="center">435.843</td>
<td valign="top" align="center">32.81</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Palemon</italic> sp.</td>
<td valign="top" align="center">3.75</td>
<td valign="top" align="center">1.02</td>
<td valign="top" align="center">4.76</td>
<td valign="top" align="center">22.7</td>
<td valign="top" align="center">1.71</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Lysmata</italic> sp.</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.38</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.58</td>
<td valign="top" align="center">0.19</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Athanas nitescens</italic>
</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.04</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.0</td>
<td valign="top" align="center">0.15</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Palaemon elegans</italic>
</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.26</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.40</td>
<td valign="top" align="center">0.18</td>
</tr>
<tr>
<td valign="middle" align="left">Dendrobranchiata</td>
<td valign="top" align="center">6.25</td>
<td valign="top" align="center">0.94</td>
<td valign="top" align="center">7.94</td>
<td valign="top" align="center">57.10</td>
<td valign="top" align="center">4.30</td>
</tr>
<tr>
<td valign="middle" align="left">Decapoda n.d.</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.04</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.04</td>
<td valign="top" align="center">0.15</td>
</tr>
<tr>
<td valign="middle" align="left">
<bold>Total Decapoda</bold>
</td>
<td valign="top" align="center">
<bold>61.25</bold>
</td>
<td valign="top" align="center">
<bold>42.07</bold>
</td>
<td valign="top" align="center">
<bold>63.49</bold>
</td>
<td valign="top" align="center">
<bold>1039.57</bold>
</td>
<td valign="top" align="center">
<bold>78.26</bold>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Lysianassa</italic> sp.</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.04</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.04</td>
<td valign="top" align="center">0.15</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Dexamine spinosa</italic>
</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.04</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.0</td>
<td valign="top" align="center">0.15</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Microdeutopus</italic> sp.</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.04</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.04</td>
<td valign="top" align="center">0.15</td>
</tr>
<tr>
<td valign="middle" align="left">Amphipoda</td>
<td valign="top" align="center">12.50</td>
<td valign="top" align="center">0.34</td>
<td valign="top" align="center">11.11</td>
<td valign="top" align="center">142.67</td>
<td valign="top" align="center">10.74</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Dynamenella</italic> sp.</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.04</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.04</td>
<td valign="top" align="center">0.15</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Sphaeroma</italic> sp.</td>
<td valign="top" align="center">7.50</td>
<td valign="top" align="center">0.68</td>
<td valign="top" align="center">3.17</td>
<td valign="top" align="center">25.97</td>
<td valign="top" align="center">1.95</td>
</tr>
<tr>
<td valign="middle" align="left">Isopoda</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.11</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.16</td>
<td valign="top" align="center">0.16</td>
</tr>
<tr>
<td valign="middle" align="left">
<bold>Total Peracarida</bold>
</td>
<td valign="top" align="center">
<bold>26.25</bold>
</td>
<td valign="top" align="center">
<bold>1.28</bold>
</td>
<td valign="top" align="center">
<bold>22.22</bold>
</td>
<td valign="top" align="center">
<bold>178.97</bold>
</td>
<td valign="top" align="center">
<bold>13.47</bold>
</td>
</tr>
<tr>
<td valign="middle" align="left">Crustacea n.d</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.08</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.10</td>
<td valign="top" align="center">0.16</td>
</tr>
<tr>
<td valign="middle" align="left">
<bold>Total Crustacea</bold>
</td>
<td valign="top" align="center">
<bold>88.75</bold>
</td>
<td valign="top" align="center">
<bold>43.43</bold>
</td>
<td valign="top" align="center">
<bold>87.30</bold>
</td>
<td valign="top" align="center">
<bold>1220.65</bold>
</td>
<td valign="top" align="center">
<bold>91.90</bold>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chelon auratus</italic>
</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">25.83</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">42.99</td>
<td valign="top" align="center">3.24</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Tripterygion delaisi</italic>
</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.08</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.10</td>
<td valign="top" align="center">0.16</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Gobius</italic> sp.</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">4.34</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">8.88</td>
<td valign="top" align="center">0.67</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Gobius incognitus</italic>
</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">6.34</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">12.05</td>
<td valign="top" align="center">0.91</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Thalassoma pavo</italic>
</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">18.50</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">31.36</td>
<td valign="top" align="center">2.36</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Parablennius</italic> sp.</td>
<td valign="top" align="center">2.50</td>
<td valign="top" align="center">0.83</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">5.29</td>
<td valign="top" align="center">0.40</td>
</tr>
<tr>
<td valign="middle" align="left">Osteichtyes n.d.</td>
<td valign="top" align="center">1.25</td>
<td valign="top" align="center">0.60</td>
<td valign="top" align="center">1.59</td>
<td valign="top" align="center">2.94</td>
<td valign="top" align="center">0.22</td>
</tr>
<tr>
<td valign="middle" align="left">
<bold>Total Osteichtyes</bold>
</td>
<td valign="top" align="center">
<bold>10.00</bold>
</td>
<td valign="top" align="center">
<bold>56.53</bold>
</td>
<td valign="top" align="center">
<bold>11.11</bold>
</td>
<td valign="top" align="center">
<bold>105.61</bold>
</td>
<td valign="top" align="center">
<bold>7.95</bold>
</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<p>Data provided by the present work indicated a geographical variability for some biological features of <italic>S. porcus</italic> inhabiting two different biogeographic districts of the Mediterranean Sea: the Central Mediterranean Sea (Strait of Messina) and the Adriatic Sea (Split area) (<xref ref-type="bibr" rid="B123">Span&#xf2; and De Domenico, 2017</xref>). The relevant differences mainly involved the <italic>sagittae</italic> features of the black scorpionfish, also suggesting the reliability of otolith shape and morphometrics analyses as a useful tool for its populations&#x2019; discrimination.</p>
<p>Concerning growth and age composition, our preliminary observations indicated an estimated maximum age higher in Split (8) than in Messina (5), with the von Bertalanffy growth curves which have shown K parameters higher in Messina, and L<sub>&#x221e;</sub> parameters higher in Split. The age composition was very different, with specimens from Messina that were mainly composed of individuals belonging to the third age class (3), with length-at-age values significantly higher than those shown by those from Split. All these data indicated a faster growth rate of the studied species in the Strait of Messina, as also highlighted by the growth performance index, which was higher in Messina specimens than Split. The high degree of population separation in <italic>S. porcus</italic> was widely reported, as highlighted by the differences in growth parameters and age composition detected between literature data from different geographical areas (<xref ref-type="bibr" rid="B121">Siblot-Bout&#xe9;flika, 1976</xref>; <xref ref-type="bibr" rid="B17">Bradai and Bouain, 1988</xref>; <xref ref-type="bibr" rid="B64">Jardas and Pallaoro, 1992</xref>; <xref ref-type="bibr" rid="B71">Koca, 1997</xref>, <xref ref-type="bibr" rid="B72">2002</xref>; <xref ref-type="bibr" rid="B122">Silvestri et&#xa0;al., 2002</xref>; <xref ref-type="bibr" rid="B104">Passalacqua et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B14">Bilgin and &#xc7;elik, 2009</xref>; <xref ref-type="bibr" rid="B78">La Mesa et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B69">Kendallha et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B75">Kutsyn et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B115">Sahin et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>) (<xref ref-type="table" rid="T4">
<bold>Table&#xa0;4</bold>
</xref>). Indeed, the overall comparison of the growth studies on this species from the Mediterranean basin highlighted a variability of &#x424;&#x2019; index between 1.92 and 2.3 (excluding the extreme value of 2.71 reported by <xref ref-type="bibr" rid="B14">Bilgin and &#xc7;elik, 2009</xref>, being based on an unrealistic L&#x221e; of 140 cm). The &#x424;&#x2019; values shown by results were within the range of the previous studies, but with extreme values (1.92 and 2.26), reflecting the plasticity of <italic>S. porcus</italic> growth. The L&#x221e; values reported by results were also in line with previous literature, in which they range from 19.8 and 60.94 cm. The only parameter not in line was the t<sub>0</sub> from Messina specimens, lower than those reported by literature data from other Mediterranean geographical areas.</p>
<table-wrap id="T4" position="float">
<label>Table&#xa0;4</label>
<caption>
<p>Von Bertalanffy growth parameters and derived growth performance index (&#x424;&#x2019;) of <italic>S. porcus</italic> from literature and the present study.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">L&#x221e; (cm)</th>
<th valign="middle" align="center">K (year<sup>-1</sup>)</th>
<th valign="middle" align="center">t<sub>0</sub>
</th>
<th valign="middle" align="center">&#x424;&#x2019;</th>
<th valign="middle" align="center">Area</th>
<th valign="middle" align="center">Authors</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">140.745</td>
<td valign="middle" align="center">0.026</td>
<td valign="middle" align="center">-1.557</td>
<td valign="middle" align="center">2.711</td>
<td valign="middle" align="center">Black Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B14">Bilgin and &#xc7;elik, 2009</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">40.810</td>
<td valign="middle" align="center">0.107</td>
<td valign="middle" align="center">-2.227</td>
<td valign="middle" align="center">2.250</td>
<td valign="middle" align="center">Black Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B71">Koca, 1997</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">19.8</td>
<td valign="middle" align="center">0.35</td>
<td valign="middle" align="center">-0.48</td>
<td valign="middle" align="center">2.14</td>
<td valign="middle" align="center">Black Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B75">Kutsyn et&#xa0;al., 2019</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">40.8</td>
<td valign="middle" align="center">0.11</td>
<td valign="middle" align="center">-2.23</td>
<td valign="middle" align="center">2.26</td>
<td valign="middle" align="center">Black Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B72">Koca, 2002</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">22.15</td>
<td valign="middle" align="center">0.287</td>
<td valign="middle" align="center">-1.577</td>
<td valign="middle" align="center">2.149</td>
<td valign="middle" align="center">Black Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B115">Sahin et al., 2019</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">23.1</td>
<td valign="middle" align="center">0.16</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">1.93</td>
<td valign="middle" align="center">Gulf of Gabes</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B17">Bradai and Bouain, 1988</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">24.377</td>
<td valign="middle" align="center">0.161</td>
<td valign="middle" align="center">-1.191</td>
<td valign="middle" align="center">1.981</td>
<td valign="middle" align="center">Marmara Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B135">Unsal and Oral, 1994</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">28.200</td>
<td valign="middle" align="center">0.182</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">2.161</td>
<td valign="middle" align="center">Adriatic Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B64">Jardas and Pallaoro, 1992</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">60.94</td>
<td valign="middle" align="center">0.03</td>
<td valign="middle" align="center">-5.18</td>
<td valign="middle" align="center">2.05</td>
<td valign="middle" align="center">Adriatic Sea<break/>(Split Area)</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">41.99</td>
<td valign="middle" align="center">0.06</td>
<td valign="middle" align="center">-3.54</td>
<td valign="middle" align="center">2.02</td>
<td valign="middle" align="center">Adriatic Sea<break/>(Pag Area)</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">22.30</td>
<td valign="middle" align="center">0.23</td>
<td valign="middle" align="center">-3.43</td>
<td valign="middle" align="center">2.07</td>
<td valign="middle" align="center">Adriatic Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B78">La Mesa et&#xa0;al., 2010</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">21.80</td>
<td valign="middle" align="center">0.29</td>
<td valign="middle" align="center">-2.51</td>
<td valign="middle" align="center">2.29</td>
<td valign="middle" align="center">Adriatic Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B118">Scarcella et&#xa0;al., 2011</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">32.4</td>
<td valign="middle" align="center">0.18</td>
<td valign="middle" align="center">-0.93</td>
<td valign="middle" align="center">2.28</td>
<td valign="middle" align="center">Ligurian Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B122">Silvestri et&#xa0;al., 2002</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">29.3</td>
<td valign="middle" align="center">0.16</td>
<td valign="middle" align="center">0.97</td>
<td valign="middle" align="center">2.14</td>
<td valign="middle" align="center">Algerian Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B121">Siblot-Bout&#xe9;flika, 1976</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">23.5</td>
<td valign="middle" align="center">0.28</td>
<td valign="middle" align="center">-0.03</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">Tyrrhenian Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B104">Passalacqua et&#xa0;al., 2005</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">23.96</td>
<td valign="middle" align="center">0.68</td>
<td valign="middle" align="center">0.058</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">Central Mediterranean Sea</td>
<td valign="middle" align="center">
<xref ref-type="bibr" rid="B69">Kendallha et&#xa0;al., 2018</xref>
</td>
</tr>
<tr>
<td valign="middle" align="center">28.78</td>
<td valign="middle" align="center">0.22</td>
<td valign="middle" align="center">-0.02</td>
<td valign="middle" align="center">2.26</td>
<td valign="middle" align="center">Strait of Messina</td>
<td valign="middle" align="center">Present study</td>
</tr>
<tr>
<td valign="middle" align="center">40.59</td>
<td valign="middle" align="center">0.05</td>
<td valign="middle" align="center">-2.58</td>
<td valign="middle" align="center">1.92</td>
<td valign="middle" align="center">Adriatic Sea<break/>(Split Area)</td>
<td valign="middle" align="center">Present study</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>Despite this, it must be considered also the effect of the length composition of samples from the two studied areas, influencing the curve trajectory of the younger age classes, and, consequently, the reconstruction of the entire growth rate. Further analyses with wide length classes composition, especially regarding the Messina population, are required to confirm the differences reported by results for the growth rate in the two studied geographical areas. <italic>S. porcus</italic> is a low-range benthic species with a high site fidelity capable of exhibiting a high phenotypic heterogeneity in its biological traits, often responding to local environmental conditions (<xref ref-type="bibr" rid="B102">&#xd6;zg&#xfc;l et&#xa0;al., 2019</xref>). The Strait of Messina represents a singularity inside the Mediterranean Sea, with a unique hydrographic regime and a peculiar seawater masses chemistry (<xref ref-type="bibr" rid="B30">Cucco et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B117">Savoca et&#xa0;al., 2020</xref>). It is located at the junction area between the Ionian and Tyrrhenian Sea. The meeting of these two water masses, different in physiochemical properties, occurs in an area with an enhanced morpho-bathymetrical irregularity of the bottom, resulting in a mixing process of the water masses, with strong currents regulated by tidal phases (<xref ref-type="bibr" rid="B30">Cucco et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B87">Longhitano, 2018</xref>). This intense hydro dynamism, with the massive presence of &#x201c;upwelling&#x201d; events, and the peculiar waters&#x2019; physio chemistry, have meant that this area has become an &#x201c;Atlantic Island&#x201d; inside the Mediterranean basin (as highlighted by the oxygen concentrations, nutrients, and temperature similar to those reported for the Atlantic Ocean) and a hot spot of biodiversity, with the presence of unique biological communities (<xref ref-type="bibr" rid="B31">De Domenico, 1987</xref>; <xref ref-type="bibr" rid="B28">Cortese and De Domenico, 1990</xref>; <xref ref-type="bibr" rid="B123">Span&#xf2; and De Domenico, 2017</xref>). Otherwise, the Adriatic Sea generally shows a pronounced seasonality and an enhanced longitudinal gradient in dissolved oxygen, temperature, nutrients, salinity, and chlorophyll-a (<xref ref-type="bibr" rid="B82">Lipizer et&#xa0;al., 2014</xref>), with urbane discharges, aeolian inputs, underground waters and surface runoff which represent the main nutrients sources. The eastern Adriatic Sea, to which Split area belongs, is outflowed by the East Adriatic current, bringing Levantine Intermediate Water from the Ionian Sea (<xref ref-type="bibr" rid="B100">Orli&#x107; et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B145">Zorica et&#xa0;al., 2019</xref>), with the water column characterized by a high homogeneity during winter, becoming more stratified during Summer (<xref ref-type="bibr" rid="B139">Vilibi&#x107; and Orli&#x107;, 2001</xref>). According to literature, different oceanographic features can influence the growth of species, resulting in a faster growth in populations inhabiting cold and productive water masses, than those from less productive and warmer ones (<xref ref-type="bibr" rid="B70">Kikuchi et&#xa0;al., 2021</xref>). In addition to the oceanographic differences between the water masses, also the environmental conditions experienced by the studied groups in the two different habitats were completely different. Specimens from Messina were sampled in the tidal ponds present in the beach rock formations, with reduced depth and large variations, also daily, of both biotic and abiotic conditions, related to tidal cycles and storms (<xref ref-type="bibr" rid="B19">Capillo et&#xa0;al., 2018</xref>). This results in marked physiochemical differences between the two studied areas, with higher temperature and salinity values in tidal pools from Messina than those from Split, probably allowing for the higher growth rate and mean length at age found in specimens from Messina than those from Split. Indeed, tidal pools represent an &#x201c;extreme&#x201d; environment, and they are considered a perfect case of study to explore the behavioural, physiological and morphological adaptations of marine species to peculiar hydrographic and ecological conditions (<xref ref-type="bibr" rid="B49">Gomes et&#xa0;al., 1998</xref>; <xref ref-type="bibr" rid="B19">Capillo et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B117">Savoca et&#xa0;al., 2020</xref>). They are a shelter from the intense hydro-dynamism affecting the area, a nursery area and a perfect hunting ground for many predators, <italic>S. porcus</italic> included. Indeed, beach rock represents the only natural substrate for benthic species present at these depths (<xref ref-type="bibr" rid="B123">Span&#xf2; and De Domenico, 2017</xref>), and a shelter from the strong currents that make the coastal areas of the Strait of Messina a difficult environment for habitat exploitation (<xref ref-type="bibr" rid="B52">Gravem and Morgan, 2017</xref>). This biotope hosts very peculiar benthic communities, with substantially different species composition and richness to those found in other similar Mediterranean environments (<xref ref-type="bibr" rid="B123">Span&#xf2; and De Domenico, 2017</xref>). It also houses extended vermetid formations (protected by the European Community) with a unique arrangement, being on the substrate surface, and not in the typical trottoir formation, as in the rest of the Mediterranean Sea (<xref ref-type="bibr" rid="B60">Ingrosso et&#xa0;al., 2018</xref>). Conversely, the biotope of the coastal waters near Split is the classical environment inhabited by the studied species, and other Scorpaenids, in the Mediterranean Sea. It is characterized by depths ranging from 10 and 40 m, the presence of rocky bottoms covered by algae, sandy bottoms, and patches of <italic>P. oceanica</italic>, with a relatively high biodiversity (up to 27 different recorded fish species, belonging to Sparidae, Mullidae, Mugilidae, Centracanthidae, Labridae and Gobiidae families) (<xref ref-type="bibr" rid="B61">Institute of Oceanography and Fisheries, 2023</xref>). Specimens from Split here analyzed were sampled at an average depth of 20 m, very different from the 2-3 m depth of the tidal ponds of Messina&#x2019;s beach rock formations. According to the literature, these differences in biotic and abiotic environmental conditions could drive the geographical variability in growth shown by results between the two studied areas (<xref ref-type="bibr" rid="B9">Bacha et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B80">Lek et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>).</p>
<p>It is well known that temperature can influence the metabolism, and consequently the growth rates, of several teleost species (<xref ref-type="bibr" rid="B7">Atkinson, 1994</xref>; <xref ref-type="bibr" rid="B73">Koz&#x142;owski et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B68">Jobling, 2011</xref>), but also food availability and quality can shape them (<xref ref-type="bibr" rid="B93">Mommsen, 1998</xref>; <xref ref-type="bibr" rid="B9">Bacha et&#xa0;al., 2010</xref>). Concerning diet composition, a comparison between the data provided by the stomach content analysis of the specimens from Messina, and the literature data regarding the studied species from Split, showed a similarity between the feeding habits of the black scorpion fish in the two areas, especially regarding the major taxa of the main preys. Results from the stomach content analysis on specimens sampled in Messina showed a high selectivity for decapods&#x2019; preys, as assessed also for other Scorpaenidae species from other geographical areas (<xref ref-type="bibr" rid="B54">Harmelin-Vivien et&#xa0;al., 1989</xref>; <xref ref-type="bibr" rid="B10">Ba&#x15f;&#xe7;&#xef;nar and Sa&#x11f;lam, 2009</xref>; <xref ref-type="bibr" rid="B26">Compaire et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B8">Aydin and Mazlum, 2020</xref>; <xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>). The high relevance of brachyuran decapods (e.g., <italic>X. pilipes, P. marmoratus</italic>, Brachyura) reported by results was in line with literature data on the studied species, but some differences regarding the contribution of teleost fishes, peracarids crustaceans and molluscs were detected. The differences in feeding habits with <italic>S. porcus</italic> specimens from Split mainly consisted of the complete absence of Molluscs and Anomura from the diet of Messina&#x2019;s specimens. According to the literature, these prey items represent an important source of food for Split population, together with teleost fishes and Caridea decapods, which showed a reduced occurrence in Messina specimens (<xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>). Concerning peracarids crustaceans (e.g., Amphipoda and Isopoda), they were well represented in the stomach content from Messina, but they are very marginal in the diet composition reported in literature from Split. This variability in peracarids&#x2019; preys could be strictly related to the biocenosis present in the tidal pools of the beachrock formations of Messina, rich in Amphipods and Isopods (being the perfect habitats for them) (<xref ref-type="bibr" rid="B32">Dias, 2013</xref>). Indeed, the presence of these crustaceans in the diet of this Scorpaenidae species has been assessed also in other similar formations present in Spain (Gulf of Cadiz) (<xref ref-type="bibr" rid="B26">Compaire et&#xa0;al., 2018</xref>). The wide variability in feeding habits between groups of individuals inhabiting different geographical areas probably reflects the capability of this species to adapt its diet according to the prey&#x2019;s availability, indicating a generalist and opportunistic feeding strategy (<xref ref-type="bibr" rid="B54">Harmelin-Vivien et&#xa0;al., 1989</xref>; <xref ref-type="bibr" rid="B10">Ba&#x15f;&#xe7;&#xef;nar and Sa&#x11f;lam, 2009</xref>; <xref ref-type="bibr" rid="B113">Rafrafi-Nouira et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B26">Compaire et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B8">Aydin and Mazlum, 2020</xref>; <xref ref-type="bibr" rid="B43">Ferri and Mati&#x107;-Skoko, 2021</xref>). This is an essential feature which allowed this species to inhabit a wide range of Mediterranean areas and habitats. Further analyses on a wider temporal scale, with a larger samples&#x2019; number, representative of the different ontogenetic classes, are required to better reconstruct the feeding habits and diet composition of <italic>S. porcus</italic> specimens, during the entire year, in the tidal ponds of the Strait of Messina. This is essential to understand how much diet variations, added to the other genetic and environmental features, can allow for the geographical differences in growth rates between the two groups of individuals highlighted by results.</p>
<p>Fish growth and metabolism, such as biotic and abiotic habitat features, can also influence otoliths growth and, consequently, their morphometry and shape (<xref ref-type="bibr" rid="B53">Gr&#xf8;nkj&#xe6;r, 2016</xref>). Lifestyle, diet composition, food quality and feeding fitness have a role in otoliths&#x2019; morphology, shape, growth patterns and physiology (<xref ref-type="bibr" rid="B109">Popper and Lu, 2000</xref>; <xref ref-type="bibr" rid="B111">Popper and Schilt, 2008</xref>; <xref ref-type="bibr" rid="B119">Schulz-Mirbach et&#xa0;al., 2019</xref>). According to the literature, otoliths&#x2019; shape and morphometry are influenced by water temperature, depth (e.g., the size of otoliths increases with greater depths and warmer water masses), genetics and lifestyle of the species (e.g., epipelagic species show smaller and more elongated <italic>sagittae</italic> than benthic ones) (<xref ref-type="bibr" rid="B85">Lombarte and Lleonart, 1993</xref>; <xref ref-type="bibr" rid="B84">Lombarte and Cruz, 2007</xref>; <xref ref-type="bibr" rid="B140">Volpedo et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B86">Lombarte et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B130">Tuset et&#xa0;al., 2015</xref>). Thanks to this high sensitivity to environmental conditions, otoliths (especially <italic>sagittae</italic>) have become an essential tool in fish stock assessment and population discrimination. This was confirmed by the results of <italic>sagittae</italic> analyses. The overall morphology and shape of both studied groups of individuals were in line with data from the literature, with several differences in morphometry. Tuset et&#xa0;al (<xref ref-type="bibr" rid="B131">Tuset et&#xa0;al., 2008</xref>). reported rectangularity and circularity values, for specimens from the western and central Mediterranean Sea, different from those of both Split and Messina specimens, with a most marked rectangular overall shape of the <italic>sagittae</italic> than the studied ones; while Yedier et&#xa0;al. (<xref ref-type="bibr" rid="B142">Yedi&#x307;Er and Bostanci, 2021</xref>). reported, from the Black Sea, Aegean Sea and Sea of Marmora, different shape indices values, e.g., roundness, ellipticity and aspect ratio, with a most marked elliptical shape of the <italic>sagittae</italic> than the studied ones. This shape heterogeneity of <italic>sagittae</italic> was also evident when comparing Split and Messina specimens. As highlighted by the overall contours and the statistical analysis on the morphometrical parameters between the two groups of individuals, specimens from Split showed wider, double-picked (with an enhanced <italic>antirostrum</italic>) more circular <italic>sagittae</italic>, than those from Messina. Otherwise, these last showed more lanceolate, larger, with a most marked <italic>rostrum</italic>, <italic>sagittae</italic>, characterized by higher otolith surface values and a more elliptical than circular shape. Based on the obtained results, the morphology and morphometry of the <italic>sagittae</italic> from both the analyzed geographical areas showed peculiar features, different from those reported in the literature from other ones, highlighting the high geographical heterogeneity of the studied species&#x2019; otoliths.</p>
<p>Several factors could have induced these differences, being otoliths under a double control of genetic and environment, and sensitive to variation in physiological and metabolic individuals&#x2019; conditions. According to Vignon and Morat (<xref ref-type="bibr" rid="B138">Vignon and Morat, 2010</xref>), different environmental factors can reshape the overall <italic>sagittae</italic> outlines, while genetic variations at the intra-specific level, related to the long-time separation between the populations, only influence the shape of the otoliths locally (mainly at <italic>rostrum</italic> and <italic>antirostrum</italic> level). Concerning the environmental conditions experienced by the two groups of individuals, one of the most evident differences between the two sampling areas was the depth range, significantly higher in Split than in Messina. Indeed, it is widely reported how species and populations inhabiting deeper habitats show wider, more circular <italic>sagittae</italic> than those from shallower ones (<xref ref-type="bibr" rid="B85">Lombarte and Lleonart, 1993</xref>; <xref ref-type="bibr" rid="B94">Monteiro et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B84">Lombarte and Cruz, 2007</xref>). This finding is in line with results, which showed also a most lanceolate <italic>sagittae</italic> in specimens from Messina, inhabiting a very shallow environment, then those from Split. The soundscape can be another factor that can strongly influence the otolith morphology and development. Indeed, according to the sensory drive hypothesis postulated by Endler (<xref ref-type="bibr" rid="B40">Endler, 1992</xref>, <xref ref-type="bibr" rid="B41">1993</xref>), there is a coevolution between detected signals and sensory systems, with the speciation that may be strongly influenced by the diversification of the organisms&#x2019; sensory interactions and environment. This is strongly evident in deep species and in species that use sound to communicate (such as rockfishes) (<xref ref-type="bibr" rid="B134">Tuset et&#xa0;al., 2016b</xref>). The soundscapes experienced by individuals drive a selection on the form-function of the fishes&#x2019; auditory system, with species or groups of individuals sharing a similar surrounding soundscape that could express an inner ears&#x2019; phenotypic similarity, expressed by otolith morphology and shape (<xref ref-type="bibr" rid="B16">Bose et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B23">Chapuis et&#xa0;al., 2023</xref>). This could be also the case of the two analyzed groups of individuals of <italic>S. porcus</italic>, with the differences related to the soundscapes of the two different studied areas that may have also influenced the detected <italic>sagittae</italic> variability in morphology, morphometry and shape. Also, metabolic rate and somatic growth can influence otolith features, such as morphometry, size and shape (<xref ref-type="bibr" rid="B110">Popper et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B119">Schulz-Mirbach et&#xa0;al., 2019</xref>). Messina specimens showed faster growth than the Split ones, which (if confirmed by the further analyses required to deeply investigate the growth rates of the studied species in the two studied areas) could be related to several environmental factors, such as the most enhanced availability of food in the tidal ponds habitats (<xref ref-type="bibr" rid="B32">Dias, 2013</xref>), probably resulting in a major food intake, and the &#x201c;extreme&#x201d; environmental conditions (such as high temperature and salinity, with large seasonal values&#x2019; fluctuation) experienced by specimens from Messina. Indeed, in addition to the enhanced availability of food in tidal ponds biocenosis, another factor strongly influencing growth and, consequently, <italic>sagittae</italic> features is the water temperature. It is widely reported how teleost species show enhanced growth rates in environments characterized by higher temperatures (<xref ref-type="bibr" rid="B107">Pepin, 1991</xref>; <xref ref-type="bibr" rid="B67">Jobling, 1997</xref>). The tidal ponds are characterized by large fluctuation of temperature, with a range (between 14 and 23&#xb0; C) (<xref ref-type="bibr" rid="B117">Savoca et&#xa0;al., 2020</xref>) higher than that reported for the Split area (data reported in the literature for the first meter of the water column between 10 to 23&#xb0;C) (<xref ref-type="bibr" rid="B74">Kraus et&#xa0;al., 2019</xref>). This different range of temperature which specimens from Messina probably experienced, in addition to other factors (such as food intake), could allowed for the faster growth shown by the results. It is necessary to enlarge the sample size also to obtain more solid data from the analyses on the intra and inter-population variability of <italic>sagittae</italic>, considering the differences in specimen numbers between size classes (especially between Size classes I, totally absent in Split, and III) in both the studied areas. Indeed, data from a larger number of specimens, with a similar sample size composition between the analyzed Size Classes, can be essential to obtain an even more complete picture of the inter and intra-population variability of the studied species. Bias related to sample size has great relevance, especially in inter-population studies involving otoliths&#x2019; shape and morphometrical data, with a non-representative number of specimens, or a not-comparable number of specimens from each population, representing one of the major concerns in this research field (<xref ref-type="bibr" rid="B56">Hilborn and Walters, 1992</xref>; <xref ref-type="bibr" rid="B18">Campana and Casselman, 1993</xref>; <xref ref-type="bibr" rid="B83">Lleonart and Maynou, 2003</xref>; <xref ref-type="bibr" rid="B15">Biol&#xe9; et&#xa0;al., 2019</xref>). Future studies, involving more specimens, and combining otoliths&#x2019; shape analysis with biological or genetic analyses, are necessary to confirm and improve the data provided by results. This is essential to deepen the knowledge base regarding the inter-population differences in <italic>sagittae</italic> features and growth rates of the studied species for further stocks&#x2019; discrimination and population studies.</p>
<p>Concerning somatic growth, it is not clear its relation to metabolic rate, food intake and otoliths&#x2019; size and growth, also because faster fish growth and a most enhanced food intake do not always result in larger otoliths (<xref ref-type="bibr" rid="B57">Hoff and Fuiman, 1993</xref>; <xref ref-type="bibr" rid="B53">Gr&#xf8;nkj&#xe6;r, 2016</xref>). Concerning the geographical differences among the size classes, the statistical analyses highlighted significant shape differences in intermediate specimens (11-18 cm), but no significant differences in morphometry between specimens from the two studied areas were found in those larger than 18 cm. This could be related to the life habits of the specimens from Messina, which could pass more time inside the tidal ponds of the beach rock formations during the first part of their life, searching for shelter and good hunting ground, as reported for other teleost species in the area (<xref ref-type="bibr" rid="B19">Capillo et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B117">Savoca et&#xa0;al., 2020</xref>). Indeed, tidal pools are recognized worldwide as an important nursery area for several teleost families, that in these environments can find shelter from predators and increased preys&#x2019; availability (<xref ref-type="bibr" rid="B32">Dias, 2013</xref>; <xref ref-type="bibr" rid="B33">Dias et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B90">Mendon&#xe7;a et&#xa0;al., 2019</xref>). This could explain the more significant morphometrical differences at the geographical level between specimens smaller than 18 cm than the larger ones. Future studies comparing the otoliths belonging to specimens smaller than 10 cm from both areas can confirm this thesis, being these absent from the Split samples here analyzed. Indeed, according to the literature, otolith shape and morphometry are under the control of both genetic and environmental influences (<xref ref-type="bibr" rid="B21">Cardinale et&#xa0;al., 2004</xref>). The environment can strongly shape the sensory organs, as also highlighted by the sensory drive hypothesis, tested by Tuset et&#xa0;al. (<xref ref-type="bibr" rid="B130">Tuset et&#xa0;al., 2015</xref>) on the <italic>sagittae</italic> of others Scorpaenidae species (<italic>Sebastes</italic> spp.). Specimens smaller than 18 cm, belonging to the first and second age groups, could be more influenced by the beach rocks&#x2019; tidal ponds than the third, spending more time in these habitats. Getting larger, <italic>S. porcus</italic> individuals could, conversely, spend more time in the nearshore coastal waters (re-entering in the tidal pounds searching for food, as reported for several transient species inhabiting these environments worldwide) (<xref ref-type="bibr" rid="B32">Dias, 2013</xref>), with biotopes and depths like those present in the Split area.</p>
<p>As shown by age composition results, tidal ponds in Messina represent a nursery area for <italic>S. porcus</italic>, characterized by a relevant abundance of small-age specimens. Considering the commercial and ecological value of this species, it is required an improved conservation of this sensitive and extreme biotope. It is necessary to enhance the management practice of commercial and recreational fishing, especially in the areas near this biotope. Moreover, being this area so close to the city, it is required to preserve it against increasing urbanization, creating buffer zones between the beach rock formation and the most anthropized zones. The conservation of these ecologically relevant habitats can improve not only the abundance and distribution of <italic>S. porcus</italic>, but also those of the several species inhabiting them, preserving the biodiversity of the entire marine coastal environment.</p>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusion</title>
<p>Assess and exploring the variations in ecomorphological features and population parameters at the geographical level is essential for both conservation purposes (being morphological features, such as those related to otoliths, and population dynamics widely used in stock assessment) and ecological studies (<xref ref-type="bibr" rid="B2">Aguirre and Lombarte, 1999</xref>; <xref ref-type="bibr" rid="B86">Lombarte et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B144">Zhuang et&#xa0;al., 2015b</xref>; <xref ref-type="bibr" rid="B6">Assis et&#xa0;al., 2020</xref>). The eco-morphological adaptation of species to different habitats and environmental features is the basis of phenotypic plasticity (i.e., the capability to express different phenotypes related to different environmental factors) (<xref ref-type="bibr" rid="B137">Via et&#xa0;al., 1995</xref>). This becomes an essential concept for the identification of marine teleost populations, being assumed that each population, living under peculiar environmental conditions, can display specific phenotypes under the regulation of genetic and/or environmental mechanisms (<xref ref-type="bibr" rid="B125">Swain and Foote, 1999</xref>; <xref ref-type="bibr" rid="B51">Grabowski et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B13">Berg et&#xa0;al., 2018</xref>). This process, being also the basis of the stock&#x2019;s differentiation, assumes relevant importance in the conservation of fishery resources and the maintenance of marine ecosystems&#x2019; well-being.</p>
<p>In this regard, the present paper assessed both the reliability of <italic>S. porcus</italic> as a model species to explore the eco-morphological variability of otoliths and population dynamics plasticity, and the reliability of otoliths shape and morphometric analysis for the populations&#x2019; discrimination in the studied species. The low home-range distribution of this benthic species, its heterogeneity in growth somatic patterns and feeding habits related to different environments, added to the high geographical variability of <italic>sagittae</italic> in shape and morphometry reported by results, make <italic>S. porcus</italic> perfect to explore the influence of different environmental conditions on teleost species. Specimens inhabiting the tidal ponds of the beachrock formations could be a perfect case study to understand how teleost species can adapt to extreme environmental conditions and peculiar habitat features. Future analyses on population dynamics and seasonal distribution of the studied species in this peculiar environment are required, to understand the ecology of this species and its life habits in these habitats. Moreover, it will be also essential to provide valuable data about the species composition and ecological inter-specific dynamics existing in tidal ponds, recognized worldwide as fundamental nursery areas, and feeding grounds for several marine species, and, consequently, as important biocenosis for the well-being and the conservation of the marine biodiversity.</p>
</sec>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Material</bold>
</xref>. Further inquiries can be directed to the corresponding authors.</p>
</sec>
<sec id="s7" sec-type="ethics-statement">
<title>Ethics statement</title>
<p>Ethical approval was not required for the studies involving animals because ethical approval was not required for the study involving animals in accordance with the local legislation and institutional requirements because biological samples were obtained from landings of commercial fishing vessels and fisherman. The studies were conducted in accordance with the local legislation and institutional requirements.</p>
</sec>
<sec id="s8" sec-type="author-contributions">
<title>Author contributions</title>
<p>CD: Writing &#x2013; original draft, Visualization, Validation, Software, Investigation, Data curation, Conceptualization. SF: Writing &#x2013; review &amp; editing, Visualization, Software, Methodology, Investigation, Formal analysis, Data curation. JF: Writing &#x2013; review &amp; editing, Visualization, Validation, Supervision, Software, Methodology, Investigation, Conceptualization. MA: Writing &#x2013; review &amp; editing, Visualization, Validation, Supervision, Methodology, Investigation, Data curation, Conceptualization. NS: Writing &#x2013; review &amp; editing, Conceptualization, Validation, Supervision, Resources, Investigation, Funding acquisition. GC: Writing &#x2013; review &amp; editing, Visualization, Validation, Project administration, Methodology, Investigation, Data curation, Conceptualization. SS: Writing &#x2013; review &amp; editing, Visualization, Software, Project administration, Methodology, Investigation, Formal analysis, Data curation, Conceptualization.</p>
</sec>
</body>
<back>
<sec id="s9" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare that no financial support was received for the research, authorship, and/or publication of this article.</p>
</sec>
<sec id="s10" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s11" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s12" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fmars.2024.1347897/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fmars.2024.1347897/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="DataSheet_1.pdf" id="SM1" mimetype="application/pdf"/>
<supplementary-material xlink:href="Image_1.tif" id="SF1" mimetype="image/tiff"/>
<supplementary-material xlink:href="Image_2.tif" id="SF2" mimetype="image/tiff"/>
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