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<front>
<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>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmars.2023.1110621</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>Diatom distribution in Holocene sediments from the northern West Caroline Basin (western equatorial Pacific) and their environmental significance</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Chen</surname>
<given-names>Min</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2185250"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Huang</surname>
<given-names>Guobiao</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2116999"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Xu</surname>
<given-names>Jishang</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/847406"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wang</surname>
<given-names>Chengtao</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Xu</surname>
<given-names>Jizheng</given-names>
</name>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Qi</surname>
<given-names>Hongshuai</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2139653"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zhang</surname>
<given-names>Aimei</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
</contrib-group>    <aff id="aff1">
<sup>1</sup>
<institution>Third Institute of Oceanography, Ministry of Natural Resources</institution>, <addr-line>Xiamen</addr-line>, <country>China</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>School of Advanced Manufacturing, Fuzhou University</institution>, <addr-line>Jinjiang</addr-line>, <country>China</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Fujian Provincial Key Laboratory of Marine Physical and Geological Processes</institution>, <addr-line>Xiamen</addr-line>, <country>China</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Key Laboratory of Submarine Geosciences and Prospecting Techniques (Ministry of Education), Ocean University of China</institution>, <addr-line>Qingdao</addr-line>, <country>China</country>
</aff>
<aff id="aff5">
<sup>5</sup>
<institution>Engineering Research Center of Marine Petroleum Development and Security Safeguard (Ministry of Education), Ocean University of China</institution>, <addr-line>Qingdao</addr-line>, <country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Yanpei Zhuang, Jimei University, China</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Jinpeng Zhang, Guangzhou Marine Geological Survey, China; Chun Chen, Island Research Center of the Ministry of Natural Resources, China; Alexander Matul, P.P. Shirshov Institute of Oceanology (RAS), Russia</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Min Chen, <email xlink:href="mailto:chenmin@tio.org.cn">chenmin@tio.org.cn</email>; Jishang Xu, <email xlink:href="mailto:jishangxu@ouc.edu.cn">jishangxu@ouc.edu.cn</email>
</p>
</fn>
<fn fn-type="other" id="fn002">
<p>This article was submitted to Marine Ecosystem Ecology, a section of the journal Frontiers in Marine Science</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>09</day>
<month>02</month>
<year>2023</year>
</pub-date>
<pub-date pub-type="collection">
<year>2023</year>
</pub-date>
<volume>10</volume>
<elocation-id>1110621</elocation-id>
<history>
<date date-type="received">
<day>29</day>
<month>11</month>
<year>2022</year>
</date>
<date date-type="accepted">
<day>20</day>
<month>01</month>
<year>2023</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2023 Chen, Huang, Xu, Wang, Xu, Qi and Zhang</copyright-statement>
<copyright-year>2023</copyright-year>
<copyright-holder>Chen, Huang, Xu, Wang, Xu, Qi and Zhang</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>Diatoms are an important component of submarine biogenic sediments and often used for carrying out palaeoceanographical reconstructions. To understand the relationship between diatoms and environment in the western equatorial Pacific, diatoms from the Holocene sediments in the northern West Caroline Basin were selected for analysis. We made quantitative statistics on diatoms and divided diatom assemblages through cluster analysis. A total of 53 species or varieties of diatoms belonging to 22 genera were identified. The range of diatom abundance was 0-88,373 valves/g. The diatoms found were mainly oceanic warm-water species. Overall, <italic>Azpeitia nodulifera</italic> was the species with the highest contribution, followed by <italic>Hemidiscus cuneiformis</italic> and <italic>Thalassiosira pacifica</italic> with these three species together accounting for approximately 80% in this area. We identified four diatom groupings and divided the study area into three regions. Diatom assemblage I in the West Caroline Basin was mainly affected by the North Equatorial Counter Current (NECC), Antarctic Intermediate Water (AAIW) and upwelling, and the supply of nutrients was relatively high. Assemblage II in the West Caroline Ridge was less influenced by currents, and thus had lower abundance and species diversity than assemblage I. The complex diatom assemblages in the Yap Trench were probably controlled by Upper Circumpolar Deep Water (UCPW) and Lower Circumpolar Deep Water (LCPW).</p>
</abstract>
<kwd-group>
<kwd>diatom</kwd>
<kwd>western equatorial Pacific</kwd>
<kwd>environmental significance</kwd>
<kwd>West Caroline Basin</kwd>
<kwd>Holocene sediment</kwd>
</kwd-group>    <contract-num rid="cn001">Grant No. 2019YFE0124700</contract-num>    <contract-num rid="cn002">Grant No. GASI-02-PAC-CJ15</contract-num>    <contract-num rid="cn003">41976198, 91858203</contract-num>    <contract-sponsor id="cn001">National Key Research and Development Program of China<named-content content-type="fundref-id">10.13039/501100012166</named-content>
</contract-sponsor>    <contract-sponsor id="cn002">Ministry of Natural Resources of the People's Republic of China<named-content content-type="fundref-id">10.13039/100015809</named-content>
</contract-sponsor>    <contract-sponsor id="cn003">National Natural Science Foundation of China<named-content content-type="fundref-id">10.13039/501100001809</named-content>
</contract-sponsor>
<counts>
<fig-count count="10"/>
<table-count count="3"/>
<equation-count count="0"/>
<ref-count count="57"/>
<page-count count="12"/>
<word-count count="4883"/>
</counts>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>The West Caroline Basin is located in the core of the Western Pacific Warm Pool, where the annual average sea-surface temperature is a minimum of 28&#xb0;C (<xref ref-type="bibr" rid="B47">Wang et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B35">Qi et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B11">Chen et&#xa0;al., 2021</xref>). Sea temperature anomalies in the region influence climate change in the Asia-Pacific region and even the world by heating seawater and transporting radiant heat to the atmosphere (<xref ref-type="bibr" rid="B29">Long et&#xa0;al., 2021</xref>). Understanding the generation and burial of biological sediments in the region is crucial to understanding the global carbon cycle and changes in marine conditions (<xref ref-type="bibr" rid="B30">Lyle and Baldauf, 2015</xref>).</p>
<p>As one of the general primary producers, diatoms are an important component of submarine biogenic sediments. In sea areas where the water depth is greater than the carbonate compensation depth, fossil diatom frustules are relatively well preserved (<xref ref-type="bibr" rid="B53">Zhang et&#xa0;al., 2015</xref>). The distribution of diatoms in sediment deposits, and the factors that control their distributions, are often discussed by researchers when reconstructing both contemporary environments and paleoenvironments (<xref ref-type="bibr" rid="B9">Chen et&#xa0;al., 2019</xref>). Diatoms from upper sediments have been studied extensively in various regions (<xref ref-type="bibr" rid="B15">Esper and Gersonde, 2014</xref>; <xref ref-type="bibr" rid="B38">Ren et&#xa0;al., 2014</xref>), allowing for reliable paleoceanographic reconstructions. (<xref ref-type="bibr" rid="B45">Tsoy and Moiseenko, 2014</xref>; <xref ref-type="bibr" rid="B42">Shen et&#xa0;al., 2017b</xref>). However, only a few investigations were published in the western Pacific. In the West Philippine Basin, the relative abundance of seven tropical pelagic diatoms was greater than 20%, and their distribution pattern was influenced by the flow of the Kuroshio Current (<xref ref-type="bibr" rid="B49">Zhai et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B8">Chen et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B41">Shen et&#xa0;al., 2017a</xref>). In the southern Mariana Trench, <xref ref-type="bibr" rid="B5">Cai (2019)</xref> found a large number of <italic>Ethmodiscus rex</italic> (Rattray) Hendey and other small diatoms, dominated by warm-water and eurythermal species.</p>
<p>The purpose of this study was to investigate the species composition and abundance of diatoms in Holocene sediments of the northern West Caroline Basin. We analyzed the environmental factors that control the distribution of diatoms by considering a series of processes, including their production, sedimentation and burial, and explored the differences of diatom assemblages in different environments and their causes. This study enriched diatom research in the western Pacific and contributed to the establishment of a comprehensive diatom&#x2013;based data set, which was necessary for carrying out future paleoceanographic reconstructions in the region.</p>
</sec>
<sec id="s2">
<label>2</label>
<title>Study area</title>
<p>The study area is located in the northern West Caroline Basin (136&#xb0;0&#x2032;-142&#xb0;0&#x2032;E, 4&#xb0;0&#x2032;-9&#xb0;0&#x2032;N; <xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>). The bathymetric range of the study area is highly variable (2395-7837 m), with an average water depth of 4,015 m, including the Yap Trench (&gt; 6500&#xa0;m), the West Caroline Ridge (&lt; 3000&#xa0;m), and a flat topographic basin in the southern part (<xref ref-type="bibr" rid="B35">Qi et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B13">Chen et&#xa0;al., 2020a</xref>). In general, the water depth gradually decreases from southwest to northeast. Because the study area is located in the deep sea, it is influenced little by continental runoff and the seawater possesses a high transparency of 20-40&#xa0;m throughout the year (<xref ref-type="bibr" rid="B48">Yue et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B52">Zhang, 2020</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>
<bold>(A)</bold> Study area (dashed boxes I and II indicate the study areas of <xref ref-type="bibr" rid="B41">Shen et&#xa0;al. 2017a</xref> and <xref ref-type="bibr" rid="B5">Cai (2019)</xref>, respectively) and <bold>(B)</bold> Sampling stations. The red arrows represent surface and subsurface currents (<xref ref-type="bibr" rid="B19">Hu et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B33">Pujari et&#xa0;al., 2021</xref>), the yellow arrows represent deep currents (<xref ref-type="bibr" rid="B25">Kawano et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B26">Kender et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B13">Chen et&#xa0;al., 2020a</xref>; <xref ref-type="bibr" rid="B28">Liu et&#xa0;al., 2022</xref>), and the green arrows represent the Antarctic Intermediate Water (<xref ref-type="bibr" rid="B3">Bingham and Lukas, 1995</xref>). Abbreviations: NEC, North Equatorial Current; NECC, North Equatorial Counter Current; NEUC, North Equatorial Undercurrent; NGCC, New Guinea Coastal Current; NGCUC, New Guinea Coastal Undercurrent; LUC, Luzon Undercurrent; MC: Mindanao Current; ME, Mindanao Eddy; MUC, Mindanao Undercurrent; AAIW, Antarctic Intermediate Water; LCPW, Lower Circumpolar Deep Water; UCPW, Upper Circumpolar Deep Water.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1110621-g001.tif"/>
</fig>
<p>The West Caroline Basin experiences a typical tropical marine climate with high temperature and rainfall all year round (<xref ref-type="bibr" rid="B35">Qi et&#xa0;al., 2020</xref>). The thermocline depth in the study area is 100-150&#xa0;m. The surface seawater possesses high temperature, low salinity and low nutrient content, and is characterized by a two-layer system. A barrier layer separates the nutrient-rich deep water from the surface water (<xref ref-type="bibr" rid="B46">Vinogradov, 1981</xref>; <xref ref-type="bibr" rid="B27">Kobayashi and Takahashi, 2002</xref>; <xref ref-type="bibr" rid="B11">Chen et&#xa0;al., 2021</xref>).</p>
<p>The many currents in the western Pacific (<xref ref-type="bibr" rid="B50">Zhang, 2008</xref>) comprise water masses of different characteristics that have an important impact on phytoplankton including diatoms. Formed from the eastward diversion of Mindanao Current (MC), the North Equatorial Counter Current (NECC;1-8&#xb0;N) crosses the West Caroline Basin from west to east and influences the study area (<xref ref-type="bibr" rid="B14">Christian et&#xa0;al., 2004</xref>). The Antarctic Intermediate Water (AAIW; 500-1200 m water depth) enters the equatorial western Pacific Ocean through the Vitiaz Channel and its northwestward branch flows to and affects the study area (<xref ref-type="bibr" rid="B3">Bingham and Lukas, 1995</xref>). The deep Upper Circumpolar Deep Water (UCPW) and Lower Circumpolar Deep Water (LCPW) currents do not directly affect diatoms in the euphotic zone, but, may play a role in the sedimentation and burial of diatom frustules.</p>
</sec>
<sec id="s3">
<label>3</label>
<title>Methods</title>
<sec id="s3_1">
<label>3.1</label>
<title>Sample information</title>
<p>The studied samples were obtained from the Global Change and Air-Sea Interaction Special-West Pacific Block PAC-CJ15 Submarine Sediment and Benthic Organisms Investigation Project. The samples were collected using a box corer from April to June 2017 (<xref ref-type="bibr" rid="B55">Zhang et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B57">Zhou et&#xa0;al., 2021</xref>). We selected 80 upper sediment samples (0-10&#xa0;cm) for analysis, and the samples contained sediments since the Holocene referring to the sedimentation rate near the study area (<xref ref-type="bibr" rid="B5">Cai, 2019</xref>). The distribution of sampling stations is shown in <xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1B</bold>
</xref>.</p>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Diatom sample processing and identification</title>
<p>All samples were processed in the diatom analysis laboratory of the Third Institute of Oceanography, Ministry of Natural Resources, Xiamen, China. We prepared samples following a modified version of the method of <xref ref-type="bibr" rid="B17">Hakansson (1984)</xref>. Briefly, samples weighing ca. 5&#xa0;g were dried in an oven at 60&#xb0;C and weighed before processing. Samples were treated with 10% HCL and 30% H<sub>2</sub>O<sub>2</sub> to remove carbonates and organic material, respectively, and washed in distilled water to remove these chemicals from the solution. Samples were then soaked in distilled water for 24&#xa0;h and scattered using an ultrasonic dispersion instrument (120&#xa0;Hz) for 2&#xa0;min. Diatoms (&gt;15 &#x3bc;m) were the main components, which indicates their relationship to the environment would be reliable. Small diatoms (&lt;15 &#x3bc;m) have been shown to little effect on the results of this type of study, so were not considered (<xref ref-type="bibr" rid="B9">Chen et&#xa0;al., 2019</xref>). Therefore, samples were filtered through a 15-&#xb5;m sieve to remove microdiatoms and other material finer than 15 &#xb5;m, after which the suspension containing diatoms was concentrated to 2&#xa0;ml. When completely homogeneous, the sub-sample of the suspension was transferred to a cover slip and air-dried. Finally, three permanent slides were made for each sample using Canadian balsam as a fixative (<xref ref-type="bibr" rid="B10">Chen et&#xa0;al., 2020b</xref>). The prepared slides were observed and identified under an optical microscope (Olympus BX51, objective lens 40&#xd7;, ocular lens 20&#xd7;), and at least 300 diatom valves were counted for each sample. For incomplete individuals, valves were counted if they were more than half complete. Diatom classification and identification mainly followed illustrations from published studies (<xref ref-type="bibr" rid="B21">Jin et&#xa0;al., 1982</xref>; <xref ref-type="bibr" rid="B39">Round et&#xa0;al., 1990</xref>; <xref ref-type="bibr" rid="B22">Jin et&#xa0;al., 1991</xref>; <xref ref-type="bibr" rid="B7">Cheng et&#xa0;al., 1996</xref>; <xref ref-type="bibr" rid="B16">Guo and Qian, 2003</xref>; <xref ref-type="bibr" rid="B34">Qi and Li, 2004</xref>; <xref ref-type="bibr" rid="B43">Smol and Stoermer, 2010</xref>; <xref ref-type="bibr" rid="B6">Cheng and Gao, 2012</xref>). Because <italic>Ethmodiscus rex</italic> specimens in the samples were fragmentary, we performed only area-based qualitative statistics.</p>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Data processing and statistical methods</title>
<p>The total abundance of diatoms is the total number of valves per gram of air-dried sediment (valves/g). Species diversity was calculated using the Shannon-Weaver index (<xref ref-type="bibr" rid="B40">Shannon and Weaver, 1949</xref>). The diatom assemblages were statistically analyzed using the R software environment and correlated with various environmental factors. Prior to statistical analysis, data were first standardized for all species. In the analysis, species with &gt;2% relative abundance in at least one sample were included (<xref ref-type="bibr" rid="B20">Imbrie and Kipp, 1971</xref>), and rare species (those occurring at fewer than five stations) and data for station E13 (where no diatoms were found) were excluded. Diatom assemblages were differentiated using cluster analysis. Since detrended correspondence analysis showed a maximum gradient of 1.68 &lt; 2, redundancy analysis (RDA) was used to determine whether differences in the assemblage vectors of each sample were statistically significant.</p>
</sec>
<sec id="s3_4">
<label>3.4</label>
<title>Environmental variables</title>
<p>The environmental data obtained in this study included sea-surface temperature (SST), salinity (SSS), dissolved oxygen (DO), nutrients (silicate, phosphate and nitrate), chlorophyll a, net primary production (NPP), particulate organic carbon (POC) and water depth (WD). Data such as SST and SSS were obtained <italic>in situ</italic> using the SBE 911Plus Conductivity-Temperature-Depth system; nutrient and oxygen levels were derived from the CARS2009 dataset for 1930 to 2009 with 1/2&#xb0;-resolution (<xref ref-type="bibr" rid="B36">Qu et&#xa0;al., 2022</xref>); chlorophyll a and POC concentrations for 1997 to 2009 at 9-km resolution were downloaded from <uri xlink:href="https://oceancolor.gsfc.nasa.gov/l3/">https://oceancolor.gsfc.nasa.gov/l3/</uri>; NPP data for 1997 to 2009 with 1/6&#xb0;-resolution were downloaded from <uri xlink:href="http://www.science.oregonstate.edu/ocean.productivity/custom.php">http://www.science.oregonstate.edu/ocean.productivity/custom.php</uri>; and WD data in those figures from the GEBCO-2020 Grid at 15 arc-second intervals were downloaded from <uri xlink:href="https://data.noaa.gov/onestop">https://data.noaa.gov/onestop</uri>.</p>
</sec>
</sec>
<sec id="s4" sec-type="results">
<label>4</label>
<title>Results</title>
<sec id="s4_1">
<label>4.1</label>
<title>Environmental characteristics</title>
<p>According to the data obtained from <italic>in situ</italic> measurements and websites (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>), SST at each station varied relatively little, with a range of 28.90 to 29.27&#xb0;C (average 29.12&#xb0;C). The SSS variation was also small, from 33.94 to 34.07 (average 33.99) whereas the DO content varied from 4.52 to 4.56 ml/l (average 4.54 ml/l). Nitrate and phosphate contents varied little, ranging from 0.03 to 0.06 &#xb5;mol/l and 0.04 to 0.11 &#xb5;mol/l, respectively, whereas silicate content varied widely (1.11-2.97 &#xb5;mol/l) and showed a trend of gradual increase from the northeast to the southwest. This increase occurs because the study area is located in the main pathway for the invasion of the silica-rich AAIW into the Northwest Pacific Ocean (<xref ref-type="bibr" rid="B37">Reid, 1973</xref>; <xref ref-type="bibr" rid="B50">Zhang, 2008</xref>; <xref ref-type="bibr" rid="B54">Zhang et&#xa0;al., 2016</xref>). In addition, NPP, POC and chlorophyll a content all showed a gradual increase from north to south, indicating higher phytoplankton biomass and primary production in the southern part of the study area.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Environmental characteristics of the study area.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1110621-g002.tif"/>
</fig>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Diatom distributions</title>
<p>A total of 53 species or varieties of diatoms belonging to 22 genera were identified at 80 stations. The relative percentages of each taxa were listed in the table of supplementary material. Diatom abundance varied from 0 to 88,373 valves/g, with an average of 14,319 valves/g. Some individual stations (E13, C03, B04, A01) yielded few diatoms, although diatom abundance was generally high, especially in the southern part of the study area where the abundance was generally greater than 20,000 valves/g (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>). The diatom abundance and species diversity in the southern deep-sea basin of the study area were higher than those in the West Caroline Ridge and Yap Trench in the north. Correlation analysis showed a strong positive correlation between diatom abundance and the content of chlorophyll a and POC (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>). Silicate was the most significant environmental factor affecting the distribution of diatom abundance, and there was a negative correlation between phosphate and diatom abundance.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Total diatom abundance and Shannon-Weaver index.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1110621-g003.tif"/>
</fig>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Results of the correlation analysis between diatom abundance and environmental factors.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">PCCs</th>
<th valign="middle" align="center">WD</th>
<th valign="middle" align="center">SST</th>
<th valign="middle" align="center">SSS</th>
<th valign="middle" align="center">DO</th>
<th valign="middle" align="center">Si</th>
<th valign="middle" align="center">N</th>
<th valign="middle" align="center">P</th>
<th valign="middle" align="center">NPP</th>
<th valign="middle" align="center">Chlor</th>
<th valign="middle" align="center">POC</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">DA</td>
<td valign="middle" align="center">0.339<sup>**</sup>
</td>
<td valign="middle" align="center">0.227<sup>*</sup>
</td>
<td valign="middle" align="center">0.233<sup>*</sup>
</td>
<td valign="middle" align="center">0.224<sup>*</sup>
</td>
<td valign="middle" align="center">0.562<sup>**</sup>
</td>
<td valign="middle" align="center">0.329<sup>**</sup>
</td>
<td valign="middle" align="center">-0.367<sup>**</sup>
</td>
<td valign="middle" align="center">0.529<sup>**</sup>
</td>
<td valign="middle" align="center">0.548<sup>**</sup>
</td>
<td valign="middle" align="center">0.581<sup>**</sup>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>*indicates significant correlation (p &lt; 0.05), ** indicates significant correlation (p &lt; 0.01); WD, water depth; SST, sea-surface temperature; SSS, sea-surface salinity; DO, dissolved oxygen; Si, silicate; N, nitrate; P, phosphate; NPP, net primary production; Chlor, chlorophyll a; POC, particulate organic carbon; DA, diatom abundance.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>In this study, diatoms with relative percentages more than 10% were designated as dominant species, and those with percentages greater than 5% but less than 10% were designated as sub&#x2013;dominant species (<xref ref-type="bibr" rid="B32">Pokras and Molfino, 1986</xref>). Therefore, the dominant species were <italic>Azpeitia africanus</italic>, <italic>Azpeitia nodulifera</italic>, <italic>Coscinodiscus</italic> sp., <italic>Hemidiscus cuneiformis</italic>, <italic>Synedra</italic> sp., <italic>Synedra tabulata</italic>, <italic>Thalassiosira pacifica</italic> and <italic>Thalassiosira leptopus</italic>, with the sub-dominant species being <italic>Coscinodiscus argus</italic>, <italic>Coscinodiscus decrescens</italic>, <italic>Roperia tesselata</italic>, <italic>Thalassionema nitzschioides</italic> and <italic>Thalassiosira excentrica</italic> (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref> and <xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). <italic>Azpeitia nodulifera</italic> (an oceanic warm-water species; <xref ref-type="bibr" rid="B41">Shen et&#xa0;al., 2017a</xref>) had the highest mean percentage content, followed by <italic>H. cuneiformis</italic> (an oceanic warm-water species; <xref ref-type="bibr" rid="B41">Shen et&#xa0;al., 2017a</xref>) and <italic>T. pacifica</italic> (a cosmopolitan species; <xref ref-type="bibr" rid="B18">Halse and Syvertsen, 1996</xref>). Overall, these three species together accounted for approximately 80% in this area.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Species represented in Holocene sediments in the study area.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" colspan="3" align="left">Dominant species<break/>(Relative abundance &gt;10%)</th>
<th valign="middle" colspan="3" align="center">Sub-dominant species<break/>(Relative abundance 5-10%)</th>
</tr>
<tr>
<th valign="middle" align="left">Name</th>
<th valign="middle" align="center">Abundance range (%)</th>
<th valign="middle" align="center">Average abundance(%)</th>
<th valign="middle" align="center">Name</th>
<th valign="middle" align="center">Abundance range (%)</th>
<th valign="middle" align="center">Average abundance(%)</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">
<italic>Azpeitia africanus</italic> Janisch</td>
<td valign="middle" align="center">0-15.00</td>
<td valign="middle" align="center">2.57</td>
<td valign="middle" align="center">
<italic>Coscinodiscus argus</italic> Ehrenberg</td>
<td valign="middle" align="center">0-6.25</td>
<td valign="middle" align="center">0.84</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Azpeitia nodulifera</italic>
<break/>A. Schmidt</td>
<td valign="middle" align="center">34.19-100</td>
<td valign="middle" align="center">69.86</td>
<td valign="middle" align="center">
<italic>Coscinodiscus decrescens</italic> Grunow</td>
<td valign="middle" align="center">0-6.52</td>
<td valign="middle" align="center">0.32</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Coscinodiscus</italic> sp.</td>
<td valign="middle" align="center">0-25.00</td>
<td valign="middle" align="center">3.65</td>
<td valign="middle" align="center">
<italic>Roperia tesselata</italic> (Rop.) Grunow</td>
<td valign="middle" align="center">0-5.71</td>
<td valign="middle" align="center">0.43</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Hemidiscus cuneiformis</italic> Wallich</td>
<td valign="middle" align="center">0-19.05</td>
<td valign="middle" align="center">6.19</td>
<td valign="middle" align="center">
<italic>Thalassionema nitzschioides</italic> Grunow</td>
<td valign="middle" align="center">0-8.55</td>
<td valign="middle" align="center">0.40</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Synedra</italic> sp.</td>
<td valign="middle" align="center">0-18.43</td>
<td valign="middle" align="center">2.47</td>
<td valign="middle" align="center">
<italic>Thalassiosira excentrica</italic> (Ehr.) Cleve</td>
<td valign="middle" align="center">0-7.51</td>
<td valign="middle" align="center">0.82</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Synedra tabulata</italic>
<break/>(Ag.) Kuetzing</td>
<td valign="middle" align="center">0-29.06</td>
<td valign="middle" align="center">2.31</td>
<td valign="middle" colspan="3" rowspan="3" align="center"/>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Thalassiosira leptopus</italic> (Grun) Hasle et G. Fryxell</td>
<td valign="middle" align="center">0-25.00</td>
<td valign="middle" align="center">1.58</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Thalassiosira pacifica</italic> Gran et Angst</td>
<td valign="middle" align="center">0-11.36</td>
<td valign="middle" align="center">3.69</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<p>Station E13 (where no diatoms were found) was excluded from the calculation of the relative percentage content of diatoms in this study.</p>
</table-wrap-foot>
</table-wrap>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Distributions of dominant diatom species in Holocene sediments.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1110621-g004.tif"/>
</fig>
<p>All stations in the study area were dominated by warm-water diatoms (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>), which constituted on average 79.81% of the diatom abundance. The most abundant species were <italic>A. nodulifera</italic>, <italic>H. cuneiformis</italic> and <italic>A. africanus</italic>, which made up 78.62% of the total. Other warm-water species included <italic>Asteromphalus heptactis</italic>, <italic>Asterolampra marylandica</italic>, <italic>Asterolampra</italic> sp., <italic>Coscinodiscus radiatus</italic>, <italic>Mastogloia cocconeiformis</italic>, <italic>Rhizosolenia bergonii</italic>, <italic>Roperia tesselata</italic> and <italic>Triceratium pentacrinus</italic>. This diversity of warm-water species was consistent with the environmental conditions of the Western Pacific Warm Pool, where the study area was located. In addition, because of the location in the deep sea, the diatoms were mainly planktonic taxa (average 99.67%) and benthic diatoms occurred only sporadically in the study area (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>).</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Distribution of warm-water and benthic diatom species.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1110621-g005.tif"/>
</fig>
<p>
<italic>Ethmodiscus rex</italic> fragments were widely distributed in the West Caroline Basin (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>). The valves of <italic>E</italic>. <italic>rex</italic> are huge and readily broken, so most of the individuals recorded were present as fragments. High concentrations of <italic>E</italic>. <italic>rex</italic> fragments were mainly recorded at stations in the deeper basins and troughs in the southern part of the study area, whereas the number of fragments in the shallow West Caroline Ridge was generally low. The correlation analysis showed that <italic>E</italic>. <italic>rex</italic> abundance had a weak positive correlation with WD. The study of <italic>E</italic>. <italic>rex</italic> in the southern Mariana Trench by <xref ref-type="bibr" rid="B5">Cai (2019)</xref> also found that this species was more abundant in deeper waters and less abundant in shallower waters, and that there was a positive correlation between <italic>E</italic>. <italic>rex</italic> abundance and WD, consistent with our findings.</p>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>Distribution of <italic>Ethmodiscus rex</italic> fragments in the study area. There are definitions for 'a few' (&lt; 10,000 per g), 'many' (10,000&#x2013;1,000,000 per g) and 'a great many' (&gt; 1,000,000 per g).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1110621-g006.tif"/>
</fig>
</sec>
<sec id="s4_3">
<label>4.3</label>
<title>Statistical analysis</title>
<p>For the statistical analysis of species data, species with relative percentages &gt; 2% were selected and rare species (those occurring at fewer than five stations) were excluded. The environmental data included WD, SST, SSS, silicate, nitrate, phosphate, DO, POC, NPP and chlorophyll a. The RDA results for environmental variables and samples, and for environmental variables and diatom species, are provided in <xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref> and <xref ref-type="fig" rid="f7">
<bold>Figures&#xa0;7</bold>
</xref>, <xref ref-type="fig" rid="f8">
<bold>8</bold>
</xref>. The first two RDA axes explained 35.8% and 23.2% of the total variation, respectively. Correlation between environmental variables and RDA axes are shown by both length and angle of arrows. Using cluster analysis, the 80 samples could be divided into four groupings based on the composition of the diatom species and diatom abundance (<xref ref-type="fig" rid="f7">
<bold>Figures&#xa0;7</bold>
</xref>, <xref ref-type="fig" rid="f9">
<bold>9</bold>
</xref>).</p>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>Summary of RDA analyses.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Axes</th>
<th valign="middle" align="center">1</th>
<th valign="middle" align="center">2</th>
<th valign="middle" align="center">3</th>
<th valign="middle" align="center">4</th>
<th valign="middle" align="center">5</th>
<th valign="middle" align="center">6</th>
<th valign="middle" align="center">7</th>
<th valign="middle" align="center">8</th>
<th valign="middle" align="center">9</th>
<th valign="middle" align="center">10</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">Eigenvalues</td>
<td valign="middle" align="center">0.01033</td>
<td valign="middle" align="center">0.00669</td>
<td valign="middle" align="center">0.00384</td>
<td valign="middle" align="center">0.00274</td>
<td valign="middle" align="center">0.00169</td>
<td valign="middle" align="center">0.00104</td>
<td valign="middle" align="center">0.00092</td>
<td valign="middle" align="center">0.00088</td>
<td valign="middle" align="center">0.00051</td>
<td valign="middle" align="center">0.00020</td>
</tr>
<tr>
<td valign="middle" align="left">Proportion explained</td>
<td valign="middle" align="center">0.35827</td>
<td valign="middle" align="center">0.23220</td>
<td valign="middle" align="center">0.13328</td>
<td valign="middle" align="center">0.09490</td>
<td valign="middle" align="center">0.05856</td>
<td valign="middle" align="center">0.03594</td>
<td valign="middle" align="center">0.03185</td>
<td valign="middle" align="center">0.03065</td>
<td valign="middle" align="center">0.01754</td>
<td valign="middle" align="center">0.00679</td>
</tr>
<tr>
<td valign="middle" align="left">Cumulative proportion</td>
<td valign="middle" align="center">0.35827</td>
<td valign="middle" align="center">0.59047</td>
<td valign="middle" align="center">0.72375</td>
<td valign="middle" align="center">0.81865</td>
<td valign="middle" align="center">0.87721</td>
<td valign="middle" align="center">0.91316</td>
<td valign="middle" align="center">0.94501</td>
<td valign="middle" align="center">0.97566</td>
<td valign="middle" align="center">0.99321</td>
<td valign="middle" align="center">1.00000</td>
</tr>
</tbody>
</table>
</table-wrap>
<fig id="f7" position="float">
<label>Figure&#xa0;7</label>
<caption>
<p>RDA biplot of environmental variables and samples. The eigenvalues for RDA axis 1 and 2 are 0.010 and 0.007, respectively. SST, sea-surface temperature; SSS, sea-surface salinity; DO, dissolved oxygen; Si, silicate; P, phosphate; N, nitrate; POC, particulate organic carbon; Chlor, chlorophyll a; NPP, net primary production; WD, water depth.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1110621-g007.tif"/>
</fig>
<fig id="f8" position="float">
<label>Figure&#xa0;8</label>
<caption>
<p>RDA biplot of diatom taxa and environmental factors. The eigenvalues for RDA axis 1 and 2 are 0.010 and 0.007, respectively. SST, sea-surface temperature; SSS, sea-surface salinity; DO, dissolved oxygen; Si, silicate; P, phosphate; N, nitrate; POC, particulate organic carbon; Chlor, chlorophyll a; NPP, net primary production; WD, water depth.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1110621-g008.tif"/>
</fig>
<fig id="f9" position="float">
<label>Figure&#xa0;9</label>
<caption>
<p>Distribution of diatom assemblages in the study area. The red arrow represents the NECC, the green arrow represents the AAIW and the yellow arrows represent deep currents.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1110621-g009.tif"/>
</fig>
<sec id="s4_3_1">
<label>4.3.1</label>
<title>Diatom assemblage I: <italic>A. nodulifera</italic>-<italic>H. cuneiformis</italic>-<italic>S. tabulata</italic>-<italic>A. undulatus</italic>
</title>
<p>Assemblage I occurred at most stations, particularly in the deep-sea basins in the central and southern part of the study area. The diatoms were mainly oceanic warm-water species <italic>A. nodulifera</italic> and <italic>H. cuneiformis</italic>, with some coastal species (e.g., <italic>S. tabulata</italic>, <italic>C. argus</italic>, and <italic>Coscinodiscus divisus</italic>) and a few benthic species, such as <italic>A. undulatus</italic>. The percentage abundances of the species in the assemblage were as follows: <italic>A. nodulifera</italic> 34.19% to 90.70% (average 68.16%); <italic>H. cuneiformis</italic> 0% to 15.16% (average 5.70%); <italic>S. tabulata</italic> 0% to 29.06% (average 2.87%); and <italic>A. undulatus</italic> 0% to 2.04% (average 0.32%). The total diatom abundance in this assemblage was generally high (3,696-88,373 valves/g), with an average of 17,826 valves/g. In addition, warm-water species only accounted for 78.37% of the assemblage, because of the relatively high number of eurythermal species. Assemblage I was positively correlated with SST and silicate and negatively correlated with phosphate and represents a tropical oceanic environment.</p>
</sec>
<sec id="s4_3_2">
<label>4.3.2</label>
<title>Diatom assemblage II: <italic>A. nodulifer</italic>-<italic>H. cuneiformis</italic>-<italic>T. pacifica</italic>-<italic>C. argus</italic>
</title>
<p>Assemblage II was mainly detected in the northern part of the study area at the West Caroline Ridge. This assemblage has a species composition similar to Assemblage I, dominated by <italic>A. nodulifera</italic>, <italic>H. cuneiformis</italic> and <italic>T. pacifica</italic>. The relative percentages were: <italic>A. nodulifera</italic> 63.16% to 91.3% (average 76.89%); <italic>H. cuneiformis</italic> 0% to 19.05% (average 8.52%); <italic>T. pacifica</italic> 0% to 8.11% (average 2.41%); and the coastal species <italic>C. argus</italic> 0% to 5.41% (average 1.53%). Compared to Assemblage I, the total diatom abundance of this assemblage was markedly lower (640-2483 valves/g), with an average of 1,705 valves/g, and warm-water species accounted for 88.04% of the assemblage. Assemblage II was positively correlated with phosphate and negatively correlated with SST and silicate. The water depth of its tropical oceanic environment is shallower than Assemblage I.</p>
</sec>
<sec id="s4_3_3">
<label>4.3.3</label>
<title>Diatom assemblage III: <italic>A. nodulifera</italic>-<italic>H. cuneiformis</italic>-<italic>C. argus</italic>
</title>
<p>Assemblage III was detected at two stations, A01 and B04, located in the waters near the Yap Trench. This assemblage exhibited both a low total diatom abundance (average 132 valves/g) and a relatively homogeneous species composition, with only three diatom species identified. The average abundances of <italic>A. nodulifera</italic>, <italic>H. cuneiformis</italic> and <italic>C. argus</italic> were 87.5%, 9.375% and 3.125%, respectively. Warm-water species represented a maximum of 96.875% of the total. Assemblage III was positively correlated with phosphate and DO.</p>
</sec>
<sec id="s4_3_4">
<label>4.3.4</label>
<title>Diatom assemblage IV (station C03): <italic>A.&#xa0;nodulifera</italic>-<italic>T. leptopus</italic>
</title>
<p>The fourth assemblage only included station C03, located in the southern Yap Trench. The total abundance of diatoms at this station was only 76 valves/g, of which <italic>A. nodulifera</italic> constituted 50%, <italic>T. leptopus</italic> made up 25%, and the remainder were <italic>Coscinodiscus</italic> sp. The proportion of warm-water species was 50%. Assemblage IV was negatively correlated with DO and WD.</p>
</sec>
</sec>
</sec>
<sec id="s5" sec-type="discussion">
<label>5</label>
<title>Discussion</title>
<sec id="s5_1">
<label>5.1</label>
<title>Indication of diatoms in Holocene sediments to environment</title>
<p>In the study area, warm-water diatoms possessed an absolute advantage, with the lowest percentage abundance of these diatoms being 44.44% at station I06, indicating a tropical oceanic environment. The tropical pelagic species <italic>A. nodulifera</italic> was the most abundant diatom at each station, consistent with the view that <italic>A. nodulifera</italic> is the main component of diatom assemblages in the surface sediments of the tropical Pacific (<xref ref-type="bibr" rid="B23">Jous&#xe9; et&#xa0;al., 1971</xref>). <italic>E. rex</italic> fragments are also widely distributed in this area, corresponding with the nutrient-poor environment in the Western Pacific Warm Pool (<xref ref-type="bibr" rid="B1">Abrantes et&#xa0;al., 2007</xref>). Compared with study area I in the West Philippine Basin (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>), there are the same four dominant species, <italic>A. africanus</italic>, <italic>A. nodulifera</italic>, <italic>H. cuneiformis</italic> and <italic>T. leptopus</italic>, indicating that the two areas possess similar growth conditions for diatoms. However, the proportion of warm-water diatoms, especially <italic>A. nodulifera</italic>, in the West Caroline Basin was obviously greater than in the West Philippine Basin as a result of latitudinal differences and a more typical pelagic environment.</p>
<p>
<italic>T. nitzschioides</italic> is widely distributed at low latitudes, and is often used as an indicator of upwelling and increased production in coastal areas; however, the species is as abundant in nutrient-poor open oceans as in coastal areas, and is described as a cosmopolitan species (<xref ref-type="bibr" rid="B27">Kobayashi and Takahashi, 2002</xref>; <xref ref-type="bibr" rid="B38">Ren et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B30">Lyle and Baldauf, 2015</xref>). In this study, <italic>T. nitzschioides</italic> was mainly detected in the high-productivity area in the southern West Caroline Basin (<xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10</bold>
</xref>). Considerable upwelling (3-10&#xb0;N, strongest at 6-7&#xb0;N) is observed in this area (<xref ref-type="bibr" rid="B29">Long et&#xa0;al., 2021</xref>), suggesting that <italic>T. nitzschioides</italic> is also good proxy for upwelling and increased production in the open ocean. Similarly, <xref ref-type="bibr" rid="B27">Kobayashi and Takahashi (2002)</xref> found <italic>T. nitzschioides</italic> may reflect weak upwelling in the equatorial western Pacific.</p>
<fig id="f10" position="float">
<label>Figure&#xa0;10</label>
<caption>
<p>Distribution of <italic>T. nitzschioides</italic> in the study area.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1110621-g010.tif"/>
</fig>
</sec>
<sec id="s5_2">
<label>5.2</label>
<title>Diatom assemblages and their environmental significance</title>
<p>The distribution of diatoms in sediments is directly related to the different temperature and salinity conditions in the euphotic zone; however, diatom frustules experience dissolution in the water column, current transport and various geological processes during sinking and after deposition on the seafloor. As a result, there are many factors affecting the distribution of diatoms in sediments (<xref ref-type="bibr" rid="B56">Zhang et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B2">Astakhov et&#xa0;al., 2015</xref>). Using the diatom assemblage distribution (<xref ref-type="fig" rid="f9">
<bold>Figure&#xa0;9</bold>
</xref>), the study area was divided into three regions: West Caroline Basin, West Caroline Ridge and Yap Trench. The West Caroline Basin mainly contains diatom assemblage I, the West Caroline Ridge is dominated by diatom assemblage II, and the Yap Trench has a mixed assemblage and more complex diatom distribution.</p>
<sec id="s5_2_1">
<label>5.2.1</label>
<title>West Caroline Basin</title>
<p>Compared with the northern part of the study area, the deep-sea basin has higher primary production. Correspondingly, the total abundance of assemblage I is obviously higher than other assemblages. Sufficient nutrients are needed for diatoms to flourish in this area, and the supply of nutrients is likely from horizontal and vertical current transport.</p>
<p>Previous studies have shown that the NECC is the main cause of high chlorophyll a in the surface water of the Western Pacific Warm Pool and the New Guinea Coastal Current transports nutrients that fertilize the southern waters of the NECC region (<xref ref-type="bibr" rid="B14">Christian et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B12">Chen et&#xa0;al., 2017</xref>). In general, the west-to-east NECC delivers warm, nutrient-rich water, which is conducive to the growth and reproduction of diatoms in the study area. In addition, the AAIW exhibits low temperatures and high silicon and low phosphorus contents (<xref ref-type="bibr" rid="B4">Bostock et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B50">Zhang, 2008</xref>; <xref ref-type="bibr" rid="B51">Zhang, 2016</xref>; <xref ref-type="bibr" rid="B24">Kashino et&#xa0;al., 2020</xref>). The current brings silicon-rich water and increases the silicate content of surface water in the West Caroline Basin through upwelling, which also weakens the dissolution of siliceous fossils. Because of the strong influence of NECC, AAIW and upwelling in the West Caroline Basin, diatom assemblage I has higher abundance and species diversity than other assemblages.</p>
<p>Different diatom species have different nutrient requirements and the ratio of nutrients in water has a significant effect on diatom species diversity and abundance (<xref ref-type="bibr" rid="B31">Owen and Crossley, 1992</xref>). RDA analysis shows that <italic>A. nodulifera</italic> and <italic>H. cuneiformis</italic> are positively correlated with phosphate, <italic>S. tabulate</italic> is positively correlated with nitrate, whereas <italic>T. leptopus</italic> and <italic>T. pacifica</italic> are not sensitive to nutrient changes (<xref ref-type="fig" rid="f8">
<bold>Figure&#xa0;8</bold>
</xref>). Therefore, lower phosphate concentrations in the West Caroline Basin are responsible for the lower content of <italic>A. nodulifera</italic> and <italic>H. cuneiformis</italic> in assemblage I. There are also some coastal species such as <italic>S. tabulata</italic>, <italic>C. argus</italic> and <italic>C. divisus</italic>, and even the benthic species <italic>A. undulatus</italic> in assemblage I. We assume that the presence of these species is the result of NECC transport from the shallow coastal waters of the Palau Islands.</p>
</sec>
<sec id="s5_2_2">
<label>5.2.2</label>
<title>West Caroline Ridge</title>
<p>The low total abundance of assemblage II (640-2,483 valves/g) in the West Caroline Ridge is caused by the weak influence of NECC and AAIW in this region, resulting in limited supply of nutrients. The absence of <italic>T. nitzschioides</italic> in assemblage II also suggests that this region is hardly influenced by upwelling. Meanwhile, sediments on the ridge are easily scoured by currents (<xref ref-type="bibr" rid="B44">Sun et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B41">Shen et&#xa0;al., 2017a</xref>). The lack of nutrients (especially silicate) and upwelling, combined with an unfavorable topography for sediment accumulation, gives rise to the low abundance and species diversity of this assemblage.</p>
<p>As can be seen in <xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>, the phosphate levels were relatively high in the West Caroline Ridge, and assemblage II was positively correlated with phosphate. Accordingly, the relative abundance of <italic>A. nodulifera</italic> and <italic>H. cuneiformis</italic> was very high (85.42%). The presence of coastal species such as <italic>C. argus</italic>, <italic>Actinocyclus ehrenbergii</italic> and <italic>Bacteriastrum hyalinum</italic> in assemblage II indicates an input of terrestrial material from the northern islands.</p>
</sec>
<sec id="s5_2_3">
<label>5.2.3</label>
<title>Yap Trench</title>
<p>Similar to the West Caroline Ridge, the Yap Trench is located at the margin of the NECC and is less affected by this current, causing diatom abundance to be generally low. In addition, the diatom species composition varied greatly among stations in this area, and the distribution of diatom assemblages was complex. Previous studies have shown that UCPW and LCPW (deeper than UCPW) are the main deep water masses in the Yap Trench and form a counterclockwise circulation in the south of the trench (<xref ref-type="bibr" rid="B28">Liu et&#xa0;al., 2022</xref>). The mixing of diatom assemblages in the region is assumed to be caused by the large topographic variations and migration of diatom frustules via abyssal flow.</p>
</sec>
</sec>
</sec>
<sec id="s6" sec-type="conclusions">
<label>6</label>
<title>Conclusions</title>
<p>A total of 53 diatom species or varieties belonging to 22 genera were identified in Holocene sediments of the northern West Caroline Basin. Diatom abundance varied from 0 to 88,373 valves/g, with an average of 14,319 valves/g. The dominant diatom species were <italic>A. africanus</italic>, <italic>A. nodulifera</italic>, <italic>Coscinodiscus</italic> sp., <italic>H. cuneiformis</italic>, <italic>Synedra</italic> sp., <italic>S. tabulata</italic>, <italic>T. leptopus</italic> and <italic>T. pacifica</italic>. Warm-water diatom species were overwhelmingly dominant, with the tropical oceanic species <italic>A. nodulifera</italic> being the dominant component in the diatom assemblages. <italic>T. nitzschioides</italic> reflected the upwelling and increased production in the West Caroline Basin.</p>
<p>Four groupings of diatoms were identified by cluster analysis and the study area could be divided into three regions. Diatom assemblage I in the West Caroline Basin was mainly affected by NECC, AAIW and upwelling, and the supply of nutrients was sufficient to maintain relatively high productivity. Assemblage II in the West Caroline Ridge was less influenced by currents, and the diatom species were dominated by <italic>A. nodulifera</italic> and <italic>H. cuneiformis</italic>, with less species diversity. The complex diatom assemblages in the Yap Trench were probably controlled by the deep UCPW and LCPW currents.</p>
<p>This study is the first to investigate Holocene diatom assemblage distributions and reveal their reflection on the environment in the West Caroline Basin, providing fundamental new information for paleoceanographic reconstruction in the western Pacific based on fossil diatom records.</p>
</sec>
<sec id="s7" sec-type="data-availability">
<title>Data availability statement</title>
<p>The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.</p>
</sec>
<sec id="s8" sec-type="author-contributions">
<title>Author contributions</title>
<p>MC: conceptualization, methodology, writing - review and editing, supervision. GH: data curation, drawing, writing - original draft. JisX: resources, investigation, background information collection. CW: data curation. JizX: resources, investigation. HQ: supervision, writing - references editing. AZ: Chinese to English. All authors contributed to the article and approved the submitted version.</p>
</sec>
</body>
<back>
<sec id="s9" sec-type="funding-information">
<title>Funding</title>
<p>We acknowledge financial support from the National Key Research and Development Program of China (Grant No. 2019YFE0124700), the Global Change and Air-Sea Interaction Project granted by the Ministry of Natural Resources of China (Grant No. GASI-02-PACCJ15), and the National Natural Science Foundation of China (41976198, 91858203).</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>We are grateful to the crew and scientists on the GASI-02-PAC-CJ15 cruise, who collected the samples analyzed in this study. We thank Sev Kender, from Liwen Bianji (Edanz) (<uri xlink:href="http://www.liwenbianji.cn">www.liwenbianji.cn</uri>) for editing the English text of a draft of this manuscript.</p>
</ack>
<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.2023.1110621/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fmars.2023.1110621/full#supplementary-material</ext-link>
</p>
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