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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
<issn pub-type="epub">1664-462X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2025.1633436</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Plant Science</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Reducing greenhouse gas emissions via harvest residue management in eucalyptus afforestation on Brazilian sandy soils</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Freitas Brilhante de S&#xe3;o Jos&#xe9;</surname>
<given-names>Jackson</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Lisboa</surname>
<given-names>Bruno Britto</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Vieira</surname>
<given-names>Frederico Costa Beber</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Zanatta</surname>
<given-names>Josil&#xe9;ia Acordi</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Araujo</surname>
<given-names>Elias Frank</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Martins</surname>
<given-names>Juscilaine Gomes</given-names>
</name>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Bender</surname>
<given-names>Andressa Classer</given-names>
</name>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Carniel</surname>
<given-names>Eduardo</given-names>
</name>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Bayer</surname>
<given-names>Cimelio</given-names>
</name>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
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</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Vargas</surname>
<given-names>Luciano Kayser</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
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<aff id="aff1">
<sup>1</sup>
<institution>Department of Agricultural Research and Diagnosis, Department of Agriculture, Livestock, Sustainable Production and Irrigation of Rio Grande do Sul</institution>, <addr-line>Porto Alegre</addr-line>,&#xa0;<country>Brazil</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Universidade Federal do Pampa</institution>, <addr-line>S&#xe3;o Gabriel</addr-line>,&#xa0;<country>Brazil</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Embrapa Floretas</institution>, <addr-line>Colombo</addr-line>,&#xa0;<country>Brazil</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>CMPC, Celulose Riograndense</institution>, <addr-line>Gua&#xed;ba</addr-line>,&#xa0;<country>Brazil</country>
</aff>
<aff id="aff5">
<sup>5</sup>
<institution>Brazilian Institute of Environment and Renewable Natural Resources</institution>, <addr-line>Porto Alegre</addr-line>,&#xa0;<country>Brazil</country>
</aff>
<aff id="aff6">
<sup>6</sup>
<institution>Department of Soil Science, Faculty of Agronomy, Universidade Federal do Rio Grande do Sul</institution>, <addr-line>Porto Alegre</addr-line>,&#xa0;<country>Brazil</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Lucian Copolovici, Aurel Vlaicu University of Arad, Romania</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Lindsay Beaumont Hutley, Charles Darwin University, Australia</p>
<p>Muhammad Shahbaz Farooq, National Agricultural Research Centre (NARC), Pakistan</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Luciano Kayser Vargas, <email xlink:href="mailto:luciano-kayser@agricultura.rs.gov.br">luciano-kayser@agricultura.rs.gov.br</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>11</day>
<month>08</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>16</volume>
<elocation-id>1633436</elocation-id>
<history>
<date date-type="received">
<day>22</day>
<month>05</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>18</day>
<month>07</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Freitas Brilhante de S&#xe3;o Jos&#xe9;, Lisboa, Vieira, Zanatta, Araujo, Martins, Bender, Carniel, Bayer and Vargas.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Freitas Brilhante de S&#xe3;o Jos&#xe9;, Lisboa, Vieira, Zanatta, Araujo, Martins, Bender, Carniel, Bayer and Vargas</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<sec>
<title>Introduction</title>
<p>The greenhouse gas balance is a central theme in discussions related to forest ecosystems. In this context, the present study evaluated the impact of five eucalyptus harvest residue management systems on atmospheric C-CO<sub>2</sub> retention in soil, greenhouse gas (GHG) emissions, and the global warming potential (GWP) in <italic>Eucalyptus saligna</italic> plantations.</p>
</sec>
<sec>
<title>Methods</title>
<p>The management systems examined were: AR - all harvest residues retained on soil; NB - harvest residues kept on soil, except bark; NBr - harvest residues kept on soil, except branches; NR - all harvest residues (bark, branches, leaves) removed; NRs - all residues from the previous rotation and new plantation litter removed using shade cloth. Soil emissions of nitrous oxide (N<sub>2</sub>O) and methane (CH<sub>4</sub>) were monitored over 12 months (October 2016 to October 2017). Soil samples were collected to a depth of one meter to assess atmospheric C-CO<sub>2</sub> retention. </p>
</sec>
<sec>
<title>Results and discussion</title>
<p>Annual N<sub>2</sub>O emissions were low (0.11&#x2013;0.23 kg N-N<sub>2</sub>O ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>) and showed no clear relationship with the amount of nitrogen added through residues. The soil consistently functioned as a methane sink across all management systems, with CH<sub>4</sub> fluxes ranging from &#x2013;2.56 to &#x2013;3.91 kg C-CH<sub>4</sub> ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>. The highest rate of C-CO<sub>2</sub> retention in soil (&#x2013;5,540 kg C-CO<sub>2</sub> ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>) was observed under the AR management system, while the lowest (&#x2013;1,752 kg C-CO<sub>2</sub> ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>) occurred under the NRs system. AR management also resulted in the lowest global warming potential (&#x2013;33,946 kg C-CO<sub>2</sub> ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>), primarily due to soil C-CO<sub>2</sub> retention (15.43%) and carbon accumulation in biomass and wood products (84.57%). These findings demonstrate that retaining eucalyptus harvest residues in subtropical sandy soils, in conjunction with carbon sequestration in wood products, constitutes an effective forest management strategy for mitigating global warming.</p>
</sec>
</abstract>
<kwd-group>
<kwd>reforestation</kwd>
<kwd>carbon stock</kwd>
<kwd>nitrous oxide</kwd>
<kwd>methane</kwd>
<kwd>soil quality</kwd>
</kwd-group>
<counts>
<fig-count count="3"/>
<table-count count="1"/>
<equation-count count="4"/>
<ref-count count="79"/>
<page-count count="11"/>
<word-count count="5242"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Functional Plant Ecology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Forest ecosystems are recognized for their efficiency in fixing atmospheric CO<sub>2</sub> and storing substantial amounts of carbon (<xref ref-type="bibr" rid="B26">Lal, 2005</xref>). They also play a pivotal role in the greenhouse gas (GHG) balance, generally acting as sources of CO<sub>2</sub> and N<sub>2</sub>O while serving as sinks for CH<sub>4</sub> (<xref ref-type="bibr" rid="B68">Walkiewicz et&#xa0;al., 2025</xref>). In this context, while forest degradation and deforestation are major contributors to the rise in atmospheric GHG concentrations (<xref ref-type="bibr" rid="B66">van der Werf et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B48">Reygadas et&#xa0;al., 2023</xref>), planted forests offer a potential mitigation strategy (<xref ref-type="bibr" rid="B70">Waring et&#xa0;al., 2020</xref>). Worldwide, planted forests occupy approximately 294 million hectares across the five continents (<xref ref-type="bibr" rid="B17">FAO, 2020</xref>). Of this total, eucalyptus plantations occupy approximately 25 million hectares in tropical and subtropical regions (<xref ref-type="bibr" rid="B34">Mao et&#xa0;al., 2024</xref>), with 7.8 million of this area located in Brazil (<xref ref-type="bibr" rid="B23">IB&#xc1;, 2024</xref>).</p>
<p>In recent years, there has been a growing economic interest in using forest harvesting residues as a source of renewable energy (<xref ref-type="bibr" rid="B64">Udali et&#xa0;al., 2024</xref>). In Brazil, it is projected that roughly 6.4 million tons of wood residues are generated annually in the eucalyptus and pine harvesting processes (<xref ref-type="bibr" rid="B44">Pincelli et&#xa0;al., 2017</xref>). From this perspective, several forestry companies worldwide are adopting the whole-tree harvesting system, which collects, in addition to wood, other components such as branches, bark, and leaves to facilitate the removal of these materials from the field (<xref ref-type="bibr" rid="B41">Nieminen et&#xa0;al., 2016</xref>).</p>
<p>However, the removal of these residues in eucalyptus areas can have adverse effects on soil quality. Possible consequences include reduced soil fertility (<xref ref-type="bibr" rid="B37">Menegale et&#xa0;al., 2016</xref>), increased susceptibility to erosion (<xref ref-type="bibr" rid="B72">Wichert et&#xa0;al., 2018</xref>), negative influence on biological activity (<xref ref-type="bibr" rid="B6">Chaer and T&#xf3;tola, 2007</xref>) and reduced soil organic C stocks (<xref ref-type="bibr" rid="B50">Rocha et&#xa0;al., 2018</xref>). Removing eucalyptus harvesting residues can be even more impacting in sandy soils, with drastic decreases in soil organic C stocks and soil C retention rates (<xref ref-type="bibr" rid="B16">Epron et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B56">S&#xe3;o Jos&#xe9; et&#xa0;al., 2023</xref>).</p>
<p>A further aspect that should be considered and studied is the impact of removing eucalyptus harvest residues on GHG emissions. In crop areas, recent studies have evaluated the influence of residue management on N<sub>2</sub>O and CH<sub>4</sub> emissions (<xref ref-type="bibr" rid="B45">Pitombo et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B67">Vasconcelos et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B27">Langeroodi et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B38">Mirzaei et&#xa0;al., 2024</xref>), and such studies have generally found that maintaining crop residues contributes to reducing emissions of these two gases. In forest areas, the information is scarcer.</p>
<p>The major source of N<sub>2</sub>O emissions in agriculture is the application of nitrogen fertilizers, but the N present in plant residues also contributes substantially to the emissions (<xref ref-type="bibr" rid="B62">Syakila and Kroeze, 2011</xref>). The magnitude of this contribution depends on the chemical composition of the residue added to the soil (<xref ref-type="bibr" rid="B29">Li et&#xa0;al., 2016</xref>). Residues with a low C/N ratio increase N<sub>2</sub>O emissions (<xref ref-type="bibr" rid="B7">Chen et&#xa0;al., 2013</xref>). In contrast, residues with a high C/N ratio favor nitrogen immobilization, resulting in lower emissions (<xref ref-type="bibr" rid="B40">Muhammad et&#xa0;al., 2011</xref>).</p>
<p>Forest soils are recognized as significant CH<sub>4</sub> sinks due to the oxidation of this GHG by methanotrophic microorganisms (<xref ref-type="bibr" rid="B73">Wigley et&#xa0;al., 2024</xref>). However, factors such as soil temperature, moisture, fertilization, and residue management determine whether the soil will act as a source or sink of CH<sub>4</sub> (<xref ref-type="bibr" rid="B67">Vasconcelos et&#xa0;al., 2018</xref>). Generally, the input of organic substrates under anaerobic conditions promotes methanogenesis, resulting in high CH<sub>4</sub> emissions (<xref ref-type="bibr" rid="B76">Zhang et&#xa0;al., 2015</xref>). On the other hand, increased soil porosity facilitates the transport of CH<sub>4</sub> to methanotrophs, enhancing its oxidation and leading to lower net emissions (<xref ref-type="bibr" rid="B46">Prajapati and Jacinthe, 2014</xref>).</p>
<p>The GHG emissions can be used to calculate the global warming potential (GWP) of different eucalyptus harvest residue management. The GWP compares the warming potential of each gas to that of CO<sub>2</sub>, which is taken as a reference (<xref ref-type="bibr" rid="B5">Bayer et&#xa0;al., 2016</xref>). Specifically, CH<sub>4</sub> and N<sub>2</sub>O have a 100-year global warming potential 34 and 298 times higher than CO<sub>2</sub>, respectively (<xref ref-type="bibr" rid="B79">Zhou et&#xa0;al., 2023</xref>). Studies in subtropical regions have demonstrated the potential of reforestation to reduce GWP values (<xref ref-type="bibr" rid="B11">de Godoi et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B36">Martins et&#xa0;al., 2015</xref>); however, these studies did not consider the effects of eucalyptus harvest residue management on soil carbon stocks and GHG emissions. Therefore, this study aimed to evaluate the influence of eucalyptus harvest residue and litter management on the GHG balance in sandy soil in the Brazilian subtropics.</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>Experimental area and treatments</title>
<p>The experimental area was in the city of Barra do Ribeiro, in Rio Grande do Sul, the southernmost state of Brazil. The site lies near the coordinates 30&#xb0;23&#x2019;S and 51&#xb0;07&#x2019;W, at an altitude of approximately 30 m above sea level. The local climate is classified as humid subtropical (Cfa) according to the K&#xf6;ppen classification, with an average annual precipitation of approximately 1400 mm and no distinct dry season. The highest average monthly temperature does not exceed 25 &#xb0;C, while the lowest is around 14 &#xb0;C, with occasional light frosts. The local soil is classified as Quartzipsamment, characterized by a sandy texture, weak structure, low water storage capacity, and low cation exchange capacity (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S1</bold>
</xref>). More details about the experimental area can be found in <xref ref-type="bibr" rid="B55">S&#xe3;o Jos&#xe9; et&#xa0;al. (2020</xref>; <xref ref-type="bibr" rid="B54">2022</xref>; <xref ref-type="bibr" rid="B56">2023</xref>). The experiment was established in 2010 using <italic>Eucalyptus saligna</italic> (clone 2864). Each plot measured 30 &#xd7; 30 m and was planted with 100 trees arranged in a grid of 10 rows by 10 plants per row. For the analyses, we considered an inner subplot measuring 18 &#xd7; 18 m, consisting of 6 rows by 6 plants. The experimental design was a completely randomized block with four replicates and five treatments. The treatments involved five different eucalyptus residue management practices, described as follows:</p>
<list list-type="order">
<list-item>
<p>AR &#x2013; All forest residues were left on the soil (i.e., bark, branches, leaves, and the litter layer from the previous rotation), with only the trunk wood removed.</p>
</list-item>
<list-item>
<p>NB &#x2013; Same as AR, but the bark was also removed.</p>
</list-item>
<list-item>
<p>NBr &#x2013; Same as AR, but branches were also removed.</p>
</list-item>
<list-item>
<p>NR &#x2013; All eucalyptus residues (including bark, branches, leaves, and litter) were removed.</p>
</list-item>
<list-item>
<p>NRs &#x2013; Same as NR, but a shade net was also used to prevent litter from the new plantation from reaching the soil surface.</p>
</list-item>
</list>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>C and N input by crop residues and litter</title>
<p>The input of C and N was assessed at the beginning of the experiment. Branches, bark, and leaves from the previous crop were collected, their mass quantified, ground, and analyzed for C and N content to estimate the amounts contributed by each component. The accumulation of litter up to the sixth year of the current cultivation, as well as the addition of C and N through residue management, were estimated as described by <xref ref-type="bibr" rid="B56">S&#xe3;o Jos&#xe9; et&#xa0;al. (2023)</xref>.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Soil organic C stocks</title>
<p>Disturbed and undisturbed soil samples were collected in July 2016, in the 6th year of cultivation, to determine organic C content and soil density, respectively. Samples were collected from the following soil layers: 0&#x2013;2.5, 2.5&#x2013;5, 5&#x2013;10, 10&#x2013;20, 20&#x2013;30, 30&#x2013;50, 50&#x2013;75, and 75&#x2013;100 cm. Carbon stocks were calculated for the 0&#x2013;100 cm profile based on equivalent soil mass, using as a reference the system in which all harvest residues and litter from the current crop were removed (NRs). Annual rates of atmospheric C&#x2013;CO<sub>2</sub> retention in the soil (Mg ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>) were calculated as the ratio between the difference in soil C stocks relative to the reference system (NRs) and the duration of cultivation, as shown in the equation:</p>
<disp-formula>
<mml:math display="block" id="M1">
<mml:mtable>
<mml:mtr>
<mml:mtd>
<mml:mrow>
<mml:mtext mathvariant="italic">C</mml:mtext>
<mml:mo>&#x2212;</mml:mo>
<mml:mtext mathvariant="italic">CO</mml:mtext>
<mml:mn>2</mml:mn>
<mml:mo>&#xa0;</mml:mo>
<mml:mtext mathvariant="italic">anual</mml:mtext>
<mml:mo>&#xa0;</mml:mo>
<mml:mtext mathvariant="italic">retention</mml:mtext>
<mml:mo>&#xa0;</mml:mo>
<mml:mtext mathvariant="italic">rate</mml:mtext>
</mml:mrow>
</mml:mtd>
</mml:mtr>
<mml:mtr>
<mml:mtd>
<mml:mrow>
<mml:mo>=</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mtext mathvariant="italic">Treatment</mml:mtext>
<mml:mo>&#xa0;</mml:mo>
<mml:mtext mathvariant="italic">soil</mml:mtext>
<mml:mo>&#xa0;</mml:mo>
<mml:mtext mathvariant="italic">C</mml:mtext>
<mml:mo>&#xa0;</mml:mo>
<mml:mtext mathvariant="italic">stock</mml:mtext>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2212;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mtext mathvariant="italic">NRs</mml:mtext>
<mml:mo>&#xa0;</mml:mo>
<mml:mtext mathvariant="italic">C</mml:mtext>
<mml:mo>&#xa0;</mml:mo>
<mml:mtext mathvariant="italic">stock</mml:mtext>
</mml:mrow>
<mml:mrow>
<mml:mn>6</mml:mn>
<mml:mo>&#xa0;</mml:mo>
<mml:mtext mathvariant="italic">years</mml:mtext>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:mtd>
</mml:mtr>
</mml:mtable>
</mml:math>
</disp-formula>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>C accumulation in wood products</title>
<p>The accumulation of C in wood products (WPs) under different residue management treatments was estimated based on the forest productivity. In the 6th year of afforestation, the diameter at 1.30 meters height (DBH) was measured using a tape measure, and the total height (h) of the experimental trees was measured using a hypsometer. Forest productivity was estimated by the average annual increase (AAI, m<sup>3</sup> ha<sup>-1</sup> year<sup>-1</sup>), based on the volume obtained after six years using the volume equation with bark, using the model by <xref ref-type="bibr" rid="B28">Leite et&#xa0;al. (1995)</xref> presented below:</p>
<disp-formula>
<mml:math display="block" id="M2">
<mml:mtable>
<mml:mtr>
<mml:mtd>
<mml:mrow>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>V</mml:mi>
</mml:mstyle>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>=</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mn mathvariant="normal">0.000048</mml:mn>
<mml:mo>&#xd7;</mml:mo>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>D</mml:mi>
<mml:mi>B</mml:mi>
</mml:mstyle>
<mml:msup>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>H</mml:mi>
</mml:mstyle>
<mml:mrow>
<mml:mn mathvariant="normal">1.720483</mml:mn>
</mml:mrow>
</mml:msup>
<mml:mo>&#xd7;</mml:mo>
<mml:msup>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>h</mml:mi>
</mml:mstyle>
<mml:mrow>
<mml:mn mathvariant="normal">1.180736</mml:mn>
</mml:mrow>
</mml:msup>
<mml:mo>&#xd7;</mml:mo>
<mml:msup>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>e</mml:mi>
</mml:mstyle>
<mml:mrow>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:mo>&#x2212;</mml:mo>
<mml:mn mathvariant="normal">3.00555</mml:mn>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#xd7;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>t</mml:mi>
<mml:mi>x</mml:mi>
</mml:mstyle>
<mml:mo stretchy="false">/</mml:mo>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>D</mml:mi>
<mml:mi>B</mml:mi>
<mml:mi>H</mml:mi>
</mml:mstyle>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo>&#xa0;</mml:mo>
</mml:mrow>
</mml:msup>
</mml:mrow>
</mml:mtd>
</mml:mtr>
<mml:mtr>
<mml:mtd>
<mml:mrow>
<mml:mo>&#xd7;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>{</mml:mo>
<mml:mn mathvariant="normal">1</mml:mn>
<mml:mo>&#x2212;</mml:mo>
<mml:msup>
<mml:mrow>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:mfrac>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>d</mml:mi>
</mml:mstyle>
<mml:mrow>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>D</mml:mi>
<mml:mi>B</mml:mi>
<mml:mi>H</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
<mml:mrow>
<mml:mn mathvariant="normal">1</mml:mn>
<mml:mo>+</mml:mo>
<mml:mn mathvariant="normal">0.228531</mml:mn>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#xd7;</mml:mo>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>d</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:msup>
<mml:mo>}</mml:mo>
<mml:mo>+</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mstyle mathvariant="normal" mathsize="normal">
<mml:mi>&#x404;</mml:mi>
</mml:mstyle>
</mml:mrow>
</mml:mtd>
</mml:mtr>
</mml:mtable>
</mml:math>
</disp-formula>
<p>where DBH represents the diameter at 1.3 meters height; h the total height; tx equals to 0, for volume with shell, or 1, for volume without shell; d is the upper commercial diameter; ands &#x404;is the experimental error.</p>
<p>As for the estimation of soil C stocks, the productivity of the NRs (195 m&#xb3; ha<sup>&#x2212;1</sup>) at six years of age was used as a reference, allowing an estimate of WP contributions in the other treatments relative to this baseline. WPs have short- and medium-term potential for carbon sequestration. We assumed a basic wood density of 458 kg m<sup>-3</sup> (<xref ref-type="bibr" rid="B33">Londero et&#xa0;al., 2015</xref>) and a C content of 446.1 g kg<sup>-1</sup> of dry wood (<xref ref-type="bibr" rid="B49">Ribeiro et&#xa0;al., 2015</xref>) for the calculations.</p>
</sec>
<sec id="s2_5">
<label>2.5</label>
<title>Assessment of N<sub>2</sub>O and CH<sub>4</sub> emissions from soil</title>
<p>To evaluate N<sub>2</sub>O and CH<sub>4</sub> emissions, we used the closed static chamber method described by <xref ref-type="bibr" rid="B39">Mosier (1989)</xref>. In each plot, a metal base (0.24 m&#xb2; area) was inserted into the soil to a depth of 5 cm. A galvanized steel chamber [60 &#xd7; 40 &#xd7; 30 cm (L &#xd7; W &#xd7; H), 0.072 m&#xb3;] was placed over a gutter fitted to the base, and water was added to the gutter to seal the system (<xref ref-type="bibr" rid="B75">Zanatta et&#xa0;al., 2010</xref>). Two internal fans, a septum connected to a three-way valve, and a digital skewer thermometer were installed in the upper part of the chamber to monitor internal temperature. The fans were powered by a battery and activated for 30 seconds immediately before sampling to homogenize the air inside the chamber. Air samples were collected using a 20 mL polypropylene syringe through the septum via the three-way valve.</p>
<p>The collections were performed at intervals of approximately 21 days between October 2016 and October 2017, totaling 18 collections. The samples were collected between 09:00 and 11:00 in the morning at 0, 20, 40, and 60 minutes after closing the chamber on the base. After collection, the samples were stored in exetainers and kept in a refrigerator at 4&#xb0;C until analysis. The N<sub>2</sub>O and CH<sub>4</sub> contents in the air samples were analyzed by gas chromatography in a GC-14 Greenhouse model equipped with an electron capture detector (ECD) and flame ionization detector (FID), using N<sub>2</sub> as the carrier gas.</p>
<p>The N<sub>2</sub>O and CH<sub>4</sub> fluxes were calculated based on the following equation:</p>
<disp-formula>
<mml:math display="block" id="M3">
<mml:mrow>
<mml:mi>f</mml:mi>
<mml:mo>=</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mi>&#x394;</mml:mi>
<mml:mi>Q</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mi>&#x394;</mml:mi>
<mml:mi>t</mml:mi>
</mml:mrow>
</mml:mfrac>
<mml:mo>&#xa0;</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mi>P</mml:mi>
<mml:mi>V</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mi>R</mml:mi>
<mml:mi>T</mml:mi>
</mml:mrow>
</mml:mfrac>
<mml:mo>&#xa0;</mml:mo>
<mml:mfrac>
<mml:mi>M</mml:mi>
<mml:mi>A</mml:mi>
</mml:mfrac>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <italic>f</italic> is the gas flux (&#x3bc;g m<sup>&#x2212;2</sup> h<sup>&#x2212;1</sup>), &#x394;Q/&#x394;t is the change in gas concentration (N<sub>2</sub>O or CH<sub>4</sub>), <italic>P</italic> is the atmospheric pressure inside the chamber (assumed to be 1 atm), <italic>V</italic> is the chamber volume (m&#xb3;), <italic>R</italic> is the universal gas constant (0.08205 atm L mol<sup>&#x2212;1</sup> K<sup>&#x2212;1</sup>), <italic>T</italic> is the temperature inside the chamber (K), <italic>M</italic> is the molar mass of the gas (g mol<sup>&#x2212;1</sup>), and <italic>A</italic> is the chamber base area (m&#xb2;).</p>
<p>The gas fluxes were measured between 9:00 and 11:00 a.m., a time interval considered the most representative of daily average GHG fluxes (<xref ref-type="bibr" rid="B1">Alves et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B9">Costa et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B25">Jantalia et&#xa0;al., 2008</xref>). Annual cumulative soil emissions of N<sub>2</sub>O and CH<sub>4</sub> were calculated using the trapezoidal rule of integration based on the fluxes measured over one year.</p>
<p>The global warming potential (GWP), expressed in Mg C-CO<sub>2</sub> equivalents, was estimated based on the annual emissions of C-CO<sub>2</sub>, N-N<sub>2</sub>O, and C-CH<sub>4</sub> from the soil. Annual C-CO<sub>2</sub> emissions were derived from changes in soil organic carbon (SOC) stocks for each treatment, using the NRs treatment as a reference. The rate of carbon retention in the soil was calculated as previously described. The GWP of the eucalyptus harvest residue management systems was determined by summing the annual emissions of the three greenhouse gases (GHGs), considering their respective global warming potentials relative to CO<sub>2</sub> (N<sub>2</sub>O = 298, CH<sub>4</sub> = 34, CO<sub>2</sub> = 1), according to the following equation:</p>
<disp-formula>
<mml:math display="block" id="M4">
<mml:mtable>
<mml:mtr>
<mml:mtd>
<mml:mrow>
<mml:mi>G</mml:mi>
<mml:mi>W</mml:mi>
<mml:mi>P</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>k</mml:mi>
<mml:mi>g</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>C</mml:mi>
<mml:msub>
<mml:mi>O</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:mi>e</mml:mi>
<mml:mi>q</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>h</mml:mi>
<mml:msup>
<mml:mi>a</mml:mi>
<mml:mrow>
<mml:mo>&#x2212;</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
</mml:msup>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>y</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>a</mml:mi>
<mml:msup>
<mml:mi>r</mml:mi>
<mml:mrow>
<mml:mo>&#x2212;</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
</mml:msup>
</mml:mrow>
</mml:mtd>
</mml:mtr>
<mml:mtr>
<mml:mtd>
  <mml:mrow>
<mml:mo>=</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>N</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:mi>O</mml:mi>
<mml:mo>&#xd7;</mml:mo>
<mml:mn>298</mml:mn>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo>+</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:mi>C</mml:mi>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mn>4</mml:mn>
</mml:msub>
<mml:mo>&#xd7;</mml:mo>
<mml:mn>34</mml:mn>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo>+</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:mtext>&#x394;</mml:mtext>
<mml:mi>C</mml:mi>
<mml:msub>
<mml:mi>O</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo>+</mml:mo>
<mml:mi>W</mml:mi>
<mml:mi>P</mml:mi>
<mml:mi>s</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>C</mml:mi>
</mml:mrow>
</mml:mtd>
</mml:mtr>
</mml:mtable>
</mml:math>
</disp-formula>
<p>where GWP is the global warming potential; N<sub>2</sub>O and CH<sub>4</sub> represent the annual emissions of N<sub>2</sub>O and CH<sub>4</sub> from the soil in the respective harvest residue management systems, multiplied by their respective global warming potentials relative to CO<sub>2</sub>, considering a time horizon of 100 years (<xref ref-type="bibr" rid="B24">IPCC, 2006</xref>); &#x394;CO<sub>2</sub> is the variation in soil C stocks in the other treatments in comparison with NRs, which was assumed to have remained similar to the stock before afforestation; WPs C is the amount of C stored in wood products.</p>
</sec>
<sec id="s2_6">
<label>2.6</label>
<title>Soil and weather parameters</title>
<p>Simultaneously with the gas flux assessments, we collected soil samples from the 0&#x2013;10 cm layer to monitor ammonium (NH<sub>4</sub>
<sup>+</sup>) and nitrate (NO<sub>3</sub>
<sup>-</sup>) levels (<xref ref-type="bibr" rid="B63">Tedesco et&#xa0;al., 1995</xref>), and water-filled pore space (WPS) (<xref ref-type="bibr" rid="B2">Anderson et&#xa0;al., 2019</xref>). Particle density was determined from disturbed soil samples collected from the 0&#x2013;10 cm layer at three random points in each treatment (<xref ref-type="bibr" rid="B15">Embrapa, 1997</xref>). Soil temperature was measured at a depth of 5 cm using a digital rod thermometer. Air temperature and rainfall data during the study period were obtained from an automatic meteorological station approximately 7 km from the experiment.</p>
</sec>
<sec id="s2_7">
<label>2.7</label>
<title>Statistical analysis</title>
<p>The variation of soil N<sub>2</sub>O and CH<sub>4</sub> fluxes was expressed through the standard error of the mean. The CH<sub>4</sub> and N<sub>2</sub>O flux data and soil parameters were correlated through Pearson&#x2019;s correlation analysis. Linear regression analyses were used to verify the relationship between the input of C and N in the different eucalyptus harvest residue management systems and annual greenhouse gas (GHG) emissions. The GWP data were subjected to analysis of variance using the MIXED procedure (<xref ref-type="bibr" rid="B57">SAS, 2014</xref>) with the means compared by Tukey&#x2019;s test at 10% significance.</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<p>Soil N<sub>2</sub>O fluxes ranged from -13.90 to 11.15 &#xb5;g N-N<sub>2</sub>O m<sup>-2</sup> h<sup>-1</sup> in the different eucalyptus harvest management systems, characterizing a low intensity of fluxes in all treatments in this sandy soil (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1a</bold>
</xref>). Since no agricultural practices occurred during the sampling period, soil N<sub>2</sub>O fluxes remained practically constant without evident influence from eucalyptus harvest residues. N<sub>2</sub>O fluxes showed a low but significant correlation with soil NH<sub>4</sub>
<sup>+</sup> contents (<xref ref-type="table" rid="T1">
<bold>Table 1</bold>
</xref>).</p>
<table-wrap id="T1" position="float">
<label>Table 1</label>
<caption>
<p>Pearson correlation between soil <inline-formula>
<mml:math display="inline" id="im1">
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:msub>
<mml:mi>O</mml:mi>
<mml:mrow>
<mml:msup>
<mml:mn>3</mml:mn>
<mml:mo>&#x2212;</mml:mo>
</mml:msup>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>and <inline-formula>
<mml:math display="inline" id="im2">
<mml:mrow>
<mml:mi>C</mml:mi>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mrow>
<mml:msup>
<mml:mn>4</mml:mn>
<mml:mo>+</mml:mo>
</mml:msup>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> fluxes, soil temperature (ST), air temperature (AT), NO&#x2083;&#x207b; and NH&#x2084;&#x207a; concentrations, and water-filled pore space (WFPS).</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">GHG</th>
<th valign="top" align="left">ST</th>
<th valign="top" align="left">AT</th>
<th valign="top" align="left">
<inline-formula>
<mml:math display="inline" id="im3">
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:msub>
<mml:mi>O</mml:mi>
<mml:mrow>
<mml:msup>
<mml:mn>3</mml:mn>
<mml:mo>&#x2212;</mml:mo>
</mml:msup>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>
</th>
<th valign="top" align="left">
<inline-formula>
<mml:math display="inline" id="im4">
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:msub>
<mml:mi>H</mml:mi>
<mml:mrow>
<mml:msup>
<mml:mn>4</mml:mn>
<mml:mo>+</mml:mo>
</mml:msup>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula>
</th>
<th valign="top" align="left">WPS</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">N<sub>2</sub>O</td>
<td valign="top" align="left">0,06<sup>ns</sup>
</td>
<td valign="top" align="left">0,09<sup>ns</sup>
</td>
<td valign="top" align="left">0,08<sup>ns</sup>
</td>
<td valign="top" align="left">0,14<sup>*</sup>
</td>
<td valign="top" align="left">0,09<sup>ns</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">CH<sub>4</sub>
</td>
<td valign="top" align="left">-0,06<sup>ns</sup>
</td>
<td valign="top" align="left">-0,09<sup>ns</sup>
</td>
<td valign="top" align="left">-0,06<sup>ns</sup>
</td>
<td valign="top" align="left">0,03<sup>ns</sup>
</td>
<td valign="top" align="left">0,06<sup>ns</sup>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>*ns: not significant; correlation coefficient (r) with p&lt;0.05.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>N<sub>2</sub>O <bold>(a)</bold> and CH<sub>4</sub> <bold>(b)</bold> fluxes from a Quartzipsamment under different eucalyptus harvest residue management practices at six years of age, in Barra do Ribeiro, Brazil, over a one-year period.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1633436-g001.tif">
<alt-text content-type="machine-generated">Line graphs showing temporal changes over a year in two panels. Panel a displays data for &#xb5;g N-N2O per square meter per hour, and panel b for &#xb5;g C-CH4 per square meter per hour. Curves represent different treatments: AR, NB, NBr, NR, and NRs. Error bars indicate variability. The x-axis represents days, and the y-axis measures emissions or absorption levels.</alt-text>
</graphic>
</fig>
<p>Soil CH<sub>4</sub> flux ranged from -104.22 to 9.33 &#xb5;g C ha<sup>-1</sup> h<sup>-1</sup>, with a strong predominance of CH<sub>4</sub> influx into the soil. As observed in N<sub>2</sub>O fluxes, there was no difference in CH<sub>4</sub> influxes between eucalyptus harvest residue management systems (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1b</bold>
</xref>). However, there was a tendency for AR and NBr management to present higher CH<sub>4</sub> oxidation rates compared to NRs in almost all evaluation periods.</p>
<p>To assess the impact of different management systems on soil C-CO<sub>2</sub> retention, we used the system in which both harvest residues and litter were removed (NRs) as the reference. Based on this, the contributions of the other management systems to SOC retention were calculated relative to the NRs system. This approach &#x2014; using the system with minimal organic input as a baseline &#x2014; has been adopted by several authors in agricultural and forest systems (<xref ref-type="bibr" rid="B11">de Godoi et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B14">dos Santos et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B13">Dietz et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B61">Souza et&#xa0;al., 2023</xref>).</p>
<p>A significant linear relationship was observed between the annual soil C-CO<sub>2</sub> retention rates and the amount of carbon added through eucalyptus harvest residues and litter (r&#xb2; = 0.81, p = 0.03) (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2a</bold>
</xref>). The highest retention rate (&#x2013;5,540 kg C-CO<sub>2</sub> ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>) was recorded under the management system that retained both the previous crop&#x2019;s residues and the current crop&#x2019;s litter (AR). In contrast, the lowest rate (&#x2013;1,752 kg C-CO<sub>2</sub> ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>) occurred under the system where only the current crop&#x2019;s litter was retained, while the previous crop&#x2019;s residues were removed (NR). Retention values similar to those in the AR system were observed in the two systems that maintained either bark or branches (NB and NBr).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Annual rates of C-CO <sub>2</sub> retention in the soil as a function of C input from eucalyptus harvest residues <bold>(a)</bold>; annual soil N-N <sub>2</sub>O emission as a function of mineral N input from eucalyptus harvest residues <bold>(b)</bold>; and annual C-CH&#x2084; influx into the soil as a function of C input from eucalyptus harvest residues <bold>(c)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1633436-g002.tif">
<alt-text content-type="machine-generated">Three graphs illustrate relationships between carbon and nitrogen inputs and greenhouse gas emissions.  a) Plot of carbon dioxide emissions versus carbon input displays a negative correlation, with equation C-CO&#x2082; = 3697.66 - 1762.49 added C, r = 0.81, p = 0.03.  b) Plot of nitrous oxide emissions versus nitrogen input shows a slight positive trend, with equation N-N&#x2082;O = 0.1000 + 0.0014 added N, r = 0.28, p = 0.35.  c) Plot of methane emissions versus carbon input indicates a weak negative relationship, with equation C-CH&#x2084; = -1.90 - 0.33 added C, r = 0.46, p = 0.20.</alt-text>
</graphic>
</fig>
<p>Annual soil N<sub>2</sub>O emissions ranged from 0.11 to 0.23 kg N-N<sub>2</sub>O ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup> and did not show a direct relationship with the amount of N added by eucalyptus harvest residues and litter (r&#xb2; = 0.28; p = 0.35) (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2b</bold>
</xref>). Regarding methane during the one-year evaluation period, the soil showed an annual uptake of -2.56 to -3.91 kg C-CH<sub>4</sub> ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>, with a weak direct correlation with the amount of C added by harvest residues and litter (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2c</bold>
</xref>).</p>
<p>All treatments demonstrated potential for mitigating global warming. The GWP values ranged from &#x2013;24,424 to &#x2013;33,946 kg CO<sub>2</sub>eq ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup> (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>). The main contributors to this atmospheric carbon sequestration were the carbon added to the soil and the carbon stored in wood products, which offer medium- and long-term sequestration potential.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Cumulative emissions and global warming potential of the soil under eucalyptus harvest residue management. Means followed by the same letter do not differ from each other according to Tukey&#x2019;s test at 10%.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1633436-g003.tif">
<alt-text content-type="machine-generated">Bar chart showing net greenhouse gas emissions in kilograms of CO2 equivalents per hectare per year. Categories NRs, NR, NBr, NB, and AR display values of -24424, -26663, -26924, -27158, and -28496 respectively. Emissions consist of black bars for Global Warming Potential (GWP) from soil CO2, N2O, and CH4, and gray bars for WPs C. The values above each bar are 0.156, -1752, -4954, -3460, and -5450. Categories are labeled b, ab, ab, ab, and a.</alt-text>
</graphic>
</fig>
<p>The management system that retained eucalyptus harvest residues (AR) showed the lowest GWP (&#x2013;33,946 kg CO<sub>2</sub>eq ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>), representing a significant reduction in greenhouse gas emissions compared to the NRs system (p&lt; 0.10). AR was followed by NBr (&#x2013;31,879 kg CO<sub>2</sub>eq ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>), NB (&#x2013;30,619 kg CO<sub>2</sub>eq ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>), and NRs (&#x2013;28,416 kg CO<sub>2</sub>eq ha<sup>&#x2212;1</sup> year<sup>&#x2212;1</sup>), with no significant differences among these three treatments.</p>
<p>In the reference system (NRs), nearly all the GWP value was attributable to carbon stored in wood. In contrast, wood carbon contributed approximately 85% of the total GWP on average in the systems with partial or complete retention of eucalyptus residues. The remaining share was associated with the influence of harvest residue management on soil organic carbon retention.</p>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<p>The N<sub>2</sub>O fluxes we observed are consistent with recent studies in eucalyptus areas in tropical Brazilian soils. <xref ref-type="bibr" rid="B10">Cuer et&#xa0;al. (2018)</xref> found values below 10 &#x3bc;g N-N<sub>2</sub>O m<sup>&#x2212;2</sup> h<sup>&#x2212;1</sup>. <xref ref-type="bibr" rid="B60">Silveira et&#xa0;al. (2022)</xref> observed that eucalyptus forests emitted up to 5.5 &#x3bc;g N-N<sub>2</sub>O m<sup>&#x2212;2</sup> h<sup>&#x2212;1</sup> during the rainy spring, but acted as a N<sub>2</sub>O sink during the dry winter periods.</p>
<p>In general, eucalyptus plantations established in sandy soils have low nitrogen availability and, consequently, low N<sub>2</sub>O emissions (<xref ref-type="bibr" rid="B32">Livesley et&#xa0;al., 2009</xref>). In addition to the low levels of N-NH<sub>4</sub>
<sup>+</sup> and N-NO<sub>3</sub>
<sup>-</sup> in the soil, we also observed low WPS values, which may have contributed to the low N<sub>2</sub>O emission (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure S1</bold>
</xref>). The WPS values were below 25%, an unfavorable condition for N<sub>2</sub>O production by denitrification, which occurs at WPS values above 60% (<xref ref-type="bibr" rid="B3">Bateman and Baggs, 2005</xref>). In sandy soils, such as the one examined in this study, oxygen diffusion rates are higher than in clay soils, avoiding anaerobic conditions for a prolonged period, a key requirement for denitrification (<xref ref-type="bibr" rid="B52">Rochette et&#xa0;al., 2008</xref>). The lack of relationship between N<sub>2</sub>O fluxes and soil N-NO<sub>3</sub>
<sup>-</sup> levels reinforces this hypothesis. On the other hand, the correlation with N-NH<sub>4</sub>
<sup>+</sup> contents suggests that the nitrification process may have contributed significantly to N<sub>2</sub>O fluxes. During nitrification, a process favored under aerobic conditions, intermediate compounds may lead to non-obligatory N<sub>2</sub>O production (<xref ref-type="bibr" rid="B77">Zhang et&#xa0;al., 2025</xref>).</p>
<p>The impact of plant residues on N<sub>2</sub>O emissions depends on the composition of these residues, especially their C/N ratio (<xref ref-type="bibr" rid="B29">Li et&#xa0;al., 2016</xref>). In general, emissions are negatively correlated with the C/N ratio, meaning that the presence of residues with a high C/N ratio promotes N immobilization, reducing net mineralization and N<sub>2</sub>O production (<xref ref-type="bibr" rid="B43">Pilegaard et&#xa0;al., 2006</xref>). In this context, our results support the idea that the input of forest residues with a C/N ratio greater than 30 has a low contribution to N<sub>2</sub>O emissions (<xref ref-type="bibr" rid="B19">Fest et&#xa0;al., 2015</xref>) due to the rapid immobilization and limited availability of N for nitrification and denitrification processes (<xref ref-type="bibr" rid="B32">Livesley et&#xa0;al., 2009</xref>). The C/N ratio values in our treatments ranged from 111.7 to 162.7, accompanied by high lignin/N ratios, 58.1 to 69.1 (<xref ref-type="bibr" rid="B56">S&#xe3;o Jos&#xe9; et&#xa0;al., 2023</xref>), which favor microbial nitrogen immobilization. Additionally, the low-organic-matter sandy soil was fertilized only during forest establishment (<xref ref-type="bibr" rid="B55">S&#xe3;o Jos&#xe9; et&#xa0;al., 2020</xref>), contributing to low nitrogen availability and, consequently, low N<sub>2</sub>O emissions.</p>
<p>However, since we did not observe differences between treatments, our results differ from other studies involving agricultural residue management (<xref ref-type="bibr" rid="B21">Gonzaga et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B35">Maris et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B67">Vasconcelos et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B47">Reeves et&#xa0;al., 2024</xref>). These differences are possibly related to the period between the application of residues and the collection of gas samples. In those studies, the highest N<sub>2</sub>O emissions occurred immediately after the addition of plant residues, with a reduction after a few months and remaining constant over time. In our study, the evaluations only occurred six years after the experiment started. Thus, we believed that N<sub>2</sub>O emissions in the management of eucalyptus harvest residues were already reduced and stabilized, and possibly the highest emissions must have occurred at the initial time of the experiment installation.</p>
<p>The negative CH<sub>4</sub> fluxes we observed are consistent with other studies conducted in eucalyptus areas (<xref ref-type="bibr" rid="B18">Fest et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B20">Fialho et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B59">Silva et&#xa0;al., 2024</xref>). In our case, the influxes were possibly favored by the sandy texture of the soil, which, regardless of the management adopted, allowed greater oxygen diffusion and, consequently, created conditions for CH<sub>4</sub> oxidation. This assumption is supported by the findings of <xref ref-type="bibr" rid="B31">Livesley et&#xa0;al. (2011)</xref> and <xref ref-type="bibr" rid="B22">Grover et&#xa0;al. (2012)</xref>, who observed similar CH<sub>4</sub> dynamics in low-nutrient sandy soils of northern Australian Eucalypt savanna woodlands.</p>
<p>Methane influxes may also be related to low WPS values that favor CH<sub>4</sub> oxidation (<xref ref-type="bibr" rid="B30">Liu et&#xa0;al., 2019</xref>). In such a condition, the improved soil porosity and gas diffusivity facilitate the transport of CH<sub>4</sub> to methanotrophic bacteria that oxidize CH<sub>4</sub> to CO<sub>2</sub>. CH<sub>4</sub> influxes are usually inversely related to soil moisture (<xref ref-type="bibr" rid="B18">Fest et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B30">Liu et&#xa0;al., 2019</xref>). Despite this, in the present study, no relationship was observed between CH<sub>4</sub> influxes and WPS, probably due to the reduced water retention capacity of the sandy soil. CH<sub>4</sub> influx is enhanced in sandy soils, where rapid drainage occurs, preventing the maintenance of high soil water contents for prolonged periods that could determine soil reduction conditions (<xref ref-type="bibr" rid="B68">Walkiewicz et&#xa0;al., 2025</xref>). Our results were similar to recent studies on agricultural residue management (<xref ref-type="bibr" rid="B71">Wegner et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B27">Langeroodi et&#xa0;al., 2019</xref>) and in harvesting and soil preparation operations in eucalyptus areas (<xref ref-type="bibr" rid="B20">Fialho et&#xa0;al., 2018</xref>). As in our study, these authors also attributed the low effect of plant residues on CH<sub>4</sub> fluxes to the small variation in soil moisture between residue managements.</p>
<p>Removing eucalyptus harvest residues has been widely considered a management that reduces C-CO<sub>2</sub> retention in the soil (<xref ref-type="bibr" rid="B50">Rocha et&#xa0;al., 2018</xref>). This practice is more relevant in soils with lower clay contents, which have a lower capacity for physical protection of soil organic matter (<xref ref-type="bibr" rid="B12">Dieckow et&#xa0;al., 2009</xref>), causing reductions in soil C compared to the maintenance of eucalyptus harvest residues (<xref ref-type="bibr" rid="B42">Oliveira et&#xa0;al., 2018</xref>). The results obtained in this study corroborate these considerations. In addition, the capacity of the soil to function as a CO<sub>2</sub> sink depends on the biomass input (<xref ref-type="bibr" rid="B8">Concei&#xe7;&#xe3;o et&#xa0;al., 2013</xref>). This dependence is evidenced by the correlation between C-CO<sub>2</sub> retention rates in the soil and the amount of C contributed by eucalyptus harvest residues and litter (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2a</bold>
</xref>), reinforcing the importance of maintaining eucalyptus harvest residues to promote C additions, mainly in sandy soils in tropical regions (<xref ref-type="bibr" rid="B16">Epron et&#xa0;al., 2015</xref>).</p>
<p>On the other hand, annual N<sub>2</sub>O emissions were not correlated with the amount of N added by plant residues. Much of the N present in the residues may have already been released in the first months of implementation of the experiment, as observed by <xref ref-type="bibr" rid="B51">Rocha et&#xa0;al. (2016)</xref>. The authors evaluated the decomposition and release of nutrients in different management systems of eucalyptus harvest residues, observing that the management system with the maintenance of all residues released approximately 130 kg ha<sup>-1</sup> of N after 300 days from the beginning of the experiment. In addition, the high C/N ratio of the bark and branches of the residues (110 and 316, respectively) and the low N content of the litter could be causing the immobilization of N by the soil microbial population.</p>
<p>Studies evaluating annual N<sub>2</sub>O emissions in reforestation areas in subtropical regions have shown variable results. The annual N<sub>2</sub>O emissions in our study were lower than those observed in areas with <italic>Acacia mearsii</italic> (0.24 &#xb1; 1.25 kg N ha<sup>-1</sup> year<sup>-1</sup>) (<xref ref-type="bibr" rid="B11">de Godoi et&#xa0;al., 2016</xref>), <italic>Acacia auriculiformis</italic> (2.3 &#xb1; 3.1 kg N ha<sup>-1</sup> year<sup>-1</sup>), and <italic>Eucalyptus urophylla</italic> (1.9 &#xb1; 2.1 kg N ha<sup>-1</sup> year<sup>-1</sup>) (<xref ref-type="bibr" rid="B78">Zhang et&#xa0;al., 2014</xref>). However, our results were quite similar to those obtained by <xref ref-type="bibr" rid="B65">van Delden et&#xa0;al. (2018)</xref>, who observed annual N<sub>2</sub>O emissions ranging from 0.08 to 0.09 kg N ha<sup>-1</sup> year<sup>-1</sup> in eucalyptus areas grown in subtropical sandy soils in Australia. Our results demonstrate the low potential for N<sub>2</sub>O emissions in the different management of eucalyptus harvest residues in this Brazilian sandy soil.</p>
<p>Despite the low linear relationship between eucalyptus harvest residue input and CH<sub>4</sub> influxes, we observed a trend of lower influx in the NRs and NR systems, which may be related to lower soil quality compared to the AR system (<xref ref-type="bibr" rid="B54">S&#xe3;o Jos&#xe9; et&#xa0;al., 2022</xref>). The loss of soil quality results in a lower capacity to oxidize CH<sub>4</sub> (<xref ref-type="bibr" rid="B4">Bayer et&#xa0;al., 2013</xref>). <xref ref-type="bibr" rid="B74">Wu et&#xa0;al. (2019)</xref> obtained similar results. The authors observed that litter removal reduced the CH<sub>4</sub> oxidation capacity by approximately 30% compared to areas that maintained litter in coniferous forests in the Chinese subtropics. This reduction was attributed to the lower abundance of methanotrophic microorganisms due to the poor availability of low-molecular-weight organic compounds caused by litter removal.</p>
<p>Carbon sequestration is considered one of the main factors controlling GWP in agricultural systems (<xref ref-type="bibr" rid="B58">Sch&#xf6;nbach et&#xa0;al., 2012</xref>), which was confirmed by our results. Furthermore, our results demonstrate that N<sub>2</sub>O and CH<sub>4</sub> fluxes made a negligible contribution to the final GWP result, as typically observed in forests (<xref ref-type="bibr" rid="B53">Saggar et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B68">Walkiewicz et&#xa0;al., 2025</xref>). Our results are similar to those obtained by <xref ref-type="bibr" rid="B76">Zhang et&#xa0;al. (2015)</xref>, who observed that the contributions of N<sub>2</sub>O and CH<sub>4</sub> to GWP were less than 3% in forest areas in the subtropical region of China. In our study, however, the contribution of these gases was even lower, not reaching 1% (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S2</bold>
</xref>).</p>
<p>Few studies have related the effects of forest management impacts with GWP, and there is no standardization in the calculations of this variable, which makes it difficult to compare results. <xref ref-type="bibr" rid="B36">Martins et&#xa0;al. (2015)</xref> observed a 76% reduction in GWP in <italic>Eucalyptus saligna</italic> areas in Australia compared to pasture areas. However, these authors did not consider the soil C retention rate and included only CH<sub>4</sub>, N<sub>2</sub>O, and CO<sub>2</sub> emissions. <xref ref-type="bibr" rid="B69">Wang et&#xa0;al. (2022)</xref> also adopted this approach, excluding soil carbon retention rates and considering only CH<sub>4</sub>, N<sub>2</sub>O, and CO<sub>2</sub> emissions in their calculations. They were among the few, if not the only, researchers to evaluate the effect of harvest residue management on GHG emissions in forest ecosystems. However, their study focused on <italic>Cunninghamia lanceolata</italic> growing in a soil type markedly different from ours, with approximately eight times higher organic matter content. <xref ref-type="bibr" rid="B11">de Godoi et&#xa0;al. (2016)</xref>, studying <italic>Acacia mearnsii</italic> areas in the Brazilian subtropics, found that wood contributed approximately 70% of the GWP value, while soil carbon retention accounted for about 30%.</p>
<p>Estimating the GWP of local agricultural and forest systems is essential for obtaining accurate and context-specific assessments of environmental impacts. To the best of our knowledge, this is the first study to evaluate the effects of eucalyptus harvest residue management on the GHG balance in a Brazilian subtropical sandy soil. In our research, the management practice that retained all eucalyptus harvest residues and litter (AR) showed significantly lower soil-associated GWP values, indicating a greater potential for carbon sequestration compared to other management strategies. These results demonstrate that maintaining eucalyptus harvest residues, besides promoting an increase in forest productivity, represents an alternative for mitigating GHG emissions in subtropical sandy soils, both due to the potential for CH<sub>4</sub> oxidation and the storage of soil organic carbon.</p>
</sec>
</body>
<back>
<sec id="s5" 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="s6" sec-type="author-contributions">
<title>Author contributions</title>
<p>JJ: Data curation, Formal analysis, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing, Conceptualization, Investigation, Methodology, Project administration, Supervision. BL: Data curation, Formal analysis, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing, Conceptualization, Methodology, Investigation, Project administration. FV: Conceptualization, Supervision, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. JZ: Formal analysis, Methodology, Supervision, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. EA: Funding acquisition, Methodology, Project administration, Resources, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. JM: Investigation, Writing &#x2013; original draft, Methodology. AB: Investigation, Methodology, Writing &#x2013; original draft. EC: Investigation, Methodology, Writing &#x2013; original draft. CB: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Supervision, Validation, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. LV: Conceptualization, Data curation, Formal analysis, Investigation, Validation, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing.</p>
</sec>
<sec id="s7" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare financial support was received for the research and/or publication of this article. This research was funded by Celulose Riograndense &#x2013; CMPC, Foundation for Research Support of Rio Grande do Sul State (Fapergs, Innovation and Technology Network of Low Carbon Agriculture and adapted to Climate Change in Rio Grande do Sul State), the National Council for Scientific and Technological Development (CNPq), Research Centre for Greenhouse Gas Innovation (RCGI), hosted by the University of S&#xe3;o Paulo (USP) and sponsored by FAPESP &#x2013; S&#xe3;o Paulo Research Foundation (2020/15230-5), and Shell Brasil.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>The authors are grateful to CMPC for enabling assessment of the field experiment and for funding.</p>
</ack>
<sec id="s8" 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>
<p>The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision.</p>
</sec>
<sec id="s9" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
</sec>
<sec id="s10" 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="s11" 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/fpls.2025.1633436/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fpls.2025.1633436/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="DataSheet1.docx" id="SM1" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document"/>
</sec>
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