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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.2025.1649473</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Plant Science</subject>
<subj-group>
<subject>Data Report</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Complete plastome of <italic>Elaeodendron glaucum</italic> (Rottb.) Pers.: genomic resources for Celastraceae systematics</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Rinandio</surname>
<given-names>Dipta Sumeru</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Husaini</surname>
<given-names>Iin Pertiwi A.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Magandhi</surname>
<given-names>Mahat</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2861539/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Martiansyah</surname>
<given-names>Irfan</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2936069/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/formal-analysis/"/>
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<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wulansari</surname>
<given-names>Tri Yuni Indah</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Hadiah</surname>
<given-names>Julisasi Tri</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Girmansyah</surname>
<given-names>Deden</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Utami</surname>
<given-names>Nanda</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Rugayah</surname>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Irsyam</surname>
<given-names>Arifin Surya Dwipa</given-names>
</name>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Irwanto</surname>
<given-names>Rina Ratnasih</given-names>
</name>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Suhatman</surname>
<given-names>Agus</given-names>
</name>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Hariri</surname>
<given-names>Muhammad Rifqi</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2885275/overview"/>
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<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
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<role content-type="https://credit.niso.org/contributor-roles/formal-analysis/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>Research Center for Ecology and Ethnobiology, National Research and Innovation Agency</institution>, <addr-line>Cibinong, Bogor</addr-line>,&#xa0;<country>Indonesia</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Research Center for Applied Botany, National Research and Innovation Agency</institution>, <addr-line>Cibinong, Bogor</addr-line>,&#xa0;<country>Indonesia</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Department of Biology, Faculty of Mathematic and Natural Sciences, Universitas Indonesia</institution>, <addr-line>Depok</addr-line>,&#xa0;<country>Indonesia</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Research Center for Biosystematics and Evolution, National Research and Innovation Agency</institution>, <addr-line>Cibinong, Bogor</addr-line>,&#xa0;<country>Indonesia</country>
</aff>
<aff id="aff5">
<sup>5</sup>
<institution>Herbarium Bandungense, School of Life Sciences and Technology, Institut Teknologi Bandung</institution>, <addr-line>Bandung</addr-line>,&#xa0;<country>Indonesia</country>
</aff>
<aff id="aff6">
<sup>6</sup>
<institution>Directorate for Scientific Management Collection, National Research and Innovation Agency</institution>, <addr-line>Cibinong, Bogor</addr-line>,&#xa0;<country>Indonesia</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/377972/overview">Yuri Shavrukov</ext-link>, Flinders University, Australia</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/2176144/overview">Leonardo Alfredo Ornella</ext-link>, Cubiqfoods SL, Spain</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/396573/overview">Hoang Dang Khoa Do</ext-link>, Nguyen Tat Thanh University, Vietnam</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/3150572/overview">Qiang Wen</ext-link>, Jiangxi Academy of Forestry, China</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Muhammad Rifqi Hariri, <email xlink:href="mailto:muhammad.rifqi.hariri@brin.go.id">muhammad.rifqi.hariri@brin.go.id</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>01</day>
<month>09</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>16</volume>
<elocation-id>1649473</elocation-id>
<history>
<date date-type="received">
<day>18</day>
<month>06</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>11</day>
<month>08</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Rinandio, Husaini, Magandhi, Martiansyah, Wulansari, Hadiah, Girmansyah, Utami, Rugayah, Irsyam, Irwanto, Suhatman and Hariri.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Rinandio, Husaini, Magandhi, Martiansyah, Wulansari, Hadiah, Girmansyah, Utami, Rugayah, Irsyam, Irwanto, Suhatman and Hariri</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>
<kwd-group>
<kwd>Celastrales</kwd>
<kwd>gene annotation</kwd>
<kwd>genome structure</kwd>
<kwd>illumina</kwd>
<kwd>plastome assembly</kwd>
</kwd-group>
<counts>
<fig-count count="1"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="23"/>
<page-count count="5"/>
<word-count count="1657"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Functional and Applied Plant Genomics</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>
<italic>Elaeodendron glaucum</italic> (Rottb.) Pers., a woody species in the family Celastraceae, is widely distributed across Sri Lanka to South East Asia <xref ref-type="bibr" rid="B11">Hou, 1960</xref>; (<xref ref-type="bibr" rid="B17">POWO, 2025</xref>). Despite its broad geographic range and presumed ecological adaptability, genomic data for this species remain scarce (<xref ref-type="bibr" rid="B19">Simmons et&#xa0;al., 2012</xref>). In recent years, chloroplast genome characterization has emerged as a powerful tool for resolving phylogenetic relationships and elucidating evolutionary patterns in angiosperms (<xref ref-type="bibr" rid="B23">Zhang and Ma, 2024</xref>). For example, a recent comparative analysis of complete chloroplast genomes from 13 species of the genus <italic>Celastrus</italic> successfully reconstructed a well-supported phylogenetic tree, clarified relationships within the genus, and confirmed that <italic>Celastrus</italic> forms a monophyletic group with <italic>Tripterygium</italic> as its closest sister lineage. The comparative genomic analysis pinpointed distinct variable regions suitable as molecular markers for species delimitation and demonstrated that <italic>C. tonkinensis</italic> Pit. and <italic>C. hindsii</italic> Benth. are conspecific (<xref ref-type="bibr" rid="B14">Liu et&#xa0;al., 2024</xref>).</p>
<p>Chloroplast genomes are particularly valuable due to their highly conserved structure, maternal inheritance, and moderate mutation rates, which make them effective molecular markers for species identification, comparative genomics, and systematic classification (<xref ref-type="bibr" rid="B6">Daniell et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B15">Nadeem et&#xa0;al., 2018</xref>). In this context, the complete sequencing and structural annotation of the <italic>E. glaucum</italic> chloroplast genome addresses a notable gap in genomic resources and offers a critical foundation for improving taxonomic resolution within the Celastraceae (<xref ref-type="bibr" rid="B4">Coughenour et&#xa0;al., 2010</xref>). This research is especially relevant given the persistent taxonomic ambiguities within the genus <italic>Elaeodendron</italic> Jacq., which have been confounded by morphological convergence and a lack of comprehensive molecular data (<xref ref-type="bibr" rid="B19">Simmons et&#xa0;al., 2012</xref>). By generating and analyzing the complete chloroplast genome of <italic>E. glaucum</italic>, the present study provides essential data to clarify phylogenetic relationships both within the genus and among closely related taxa. Comparative analyses with other chloroplast genomes may further reveal species-specific structural variations, genomic signatures, and adaptive traits, thereby contributing to a more refined and reliable classification framework for the family.</p>
<p>The chloroplast genome in most land plants is characterized by a conserved quadripartite circular structure comprising a large single-copy (LSC) region, a small single-copy (SSC) region, and two inverted repeat (IR) regions (<xref ref-type="bibr" rid="B7">Dobrogojski et&#xa0;al., 2020</xref>). However, structural exceptions such as the loss of IRs or SSCs have been reported (<xref ref-type="bibr" rid="B22">Wang et&#xa0;al., 2024</xref>). The overall size of chloroplast genomes typically ranges from approximately 19 to 217 kilobases, with IR regions spanning 20 to 26 kilobases (NCBI Organelle Genome Resources). The chloroplast proteome includes roughly 3,000 proteins involved in crucial metabolic pathways, including photosynthesis, as well as the biosynthesis of fatty acids, amino acids, nucleotides, vitamins, hormones, and secondary metabolites (<xref ref-type="bibr" rid="B7">Dobrogojski et&#xa0;al., 2020</xref>). Most of these proteins are encoded by nuclear genes, synthesized in the cytosol, and subsequently imported into the chloroplast, while a smaller fraction is encoded by the chloroplast genome itself (<xref ref-type="bibr" rid="B9">Fu et&#xa0;al., 2022</xref>).</p>
<p>Recent advancements in chloroplast genome engineering have facilitated detailed investigations into gene function, regulatory mechanisms, and targeted genome modification (<xref ref-type="bibr" rid="B1">An et&#xa0;al., 2022</xref>). These technologies are increasingly being applied to enhance photosynthetic performance, develop nutritionally improved crops, and produce high-value bioproducts (<xref ref-type="bibr" rid="B5">Daniell et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B20">Singhal et&#xa0;al., 2023</xref>). This study reports the <italic>de novo</italic> sequencing, annotation, and structural characterization of the complete chloroplast genome of <italic>E. glaucum</italic>, presenting a valuable genomic reference for future research in taxonomy, evolutionary biology, and biotechnological innovation.</p>
</sec>
<sec id="s2">
<label>2</label>
<title>Method</title>
<sec id="s2_1">
<label>2.1</label>
<title>Plant material</title>
<p>Fresh leaf material of <italic>E. glaucum</italic> was collected from a cultivated individual maintained at the Kebun Raya Bogor (Bogor Botanic Gardens), West Java, Indonesia, under accession number III.G.189. The specimen&#x2019;s original provenance is traced to Puger, East Java.</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>DNA extraction</title>
<p>Genomic DNA was extracted from young leaf tissue using a modified cetyltrimethylammonium bromide (CTAB) protocol following <xref ref-type="bibr" rid="B8">Doyle and Doyle (1987)</xref>, with optimizations implemented to enhance yield and purity. DNA quality and quantity were initially assessed using a NanoDrop 2000 spectrophotometer (Thermo Scientific), and integrity was evaluated via 1% TBE agarose gel electrophoresis. For more accurate quantification, the Qubit dsDNA High Sensitivity Assay Kit (Thermo Scientific) was used. Fragment size distribution and integrity were further validated using the Agilent 4150 TapeStation system.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Whole genome sequencing</title>
<p>High-quality genomic DNA was subsequently subjected to library preparation. The DNA was enzymatically fragmented to produce insert-sized fragments appropriate for high-throughput sequencing. Following fragmentation, sequencing libraries were constructed and sequenced on the Illumina NextSeq 500 platform (Genetika Science Lab, Tangerang, Indonesia), generating paired-end reads of 150 base pairs. The sequencing run targeted a total yield of 10 gigabases, providing sufficient depth for comprehensive chloroplast genome assembly and downstream analyses.</p>
</sec>
<sec id="s2_4">
<label>2.3</label>
<title>Chloroplast genome assembly and annotation</title>
<p>Quality assessment of the raw sequencing reads was conducted using FastQC version 0.11.8 (<xref ref-type="bibr" rid="B2">Andrews, 2010</xref>), which provided diagnostic metrics including per-base quality scores, GC content, sequence length distribution, and indicators of potential contamination. To ensure high-fidelity reads, adapter sequences, low-quality bases (Phred score &lt;30), and nucleotide biases at the 5&#x2032; and 3&#x2032; ends were removed using Trimmomatic version 0.39 (<xref ref-type="bibr" rid="B3">Bolger et&#xa0;al., 2014</xref>). The following trimming parameters were applied: ILLUMINACLIP: TruSeq3-PE.fa:2:30:10, SLIDINGWINDOW:4:28, LEADING:28, TRAILING:28, and MINLEN:20. These processes were executed through the Galaxy web platform (<ext-link ext-link-type="uri" xlink:href="https://usegalaxy.org">https://usegalaxy.org</ext-link>, <xref ref-type="bibr" rid="B21">The Galaxy Community, 2024</xref>).</p>
<p>High-quality trimmed reads were assembled <italic>de novo</italic> into a complete chloroplast genome using GetOrganelle version 1.7.7.1 (<xref ref-type="bibr" rid="B12">Jin et&#xa0;al., 2020</xref>), an organelle-specific assembler employing a k-mer-based graph approach optimized for high-coverage plastid genomes. Genome annotation was performed using CPGAVAS2 (<xref ref-type="bibr" rid="B18">Shi et&#xa0;al., 2019</xref>) through its online platform (<ext-link ext-link-type="uri" xlink:href="http://47.96.249.172:16019/analyzer/annotate">http://47.96.249.172:16019/analyzer/annotate</ext-link>), with the chloroplast genome of <italic>Euonymus kiautschovicus</italic> Loes. (syn. <italic>Euonymus fortunei</italic> var. <italic>fortunei</italic>, GenBank accession: PQ397793) serving as the reference to guide gene prediction and structural feature identification.</p>
<p>To ensure annotation precision, subsequent manual curation and validation of coding sequences, intron-exon boundaries, and RNA genes were carried out using Unipro UGENE version 45.1 (<xref ref-type="bibr" rid="B16">Okonechnikov et&#xa0;al., 2012</xref>) and NCBI Genome Workbench version 3.8.2 (<xref ref-type="bibr" rid="B13">Kuznetsov and Bollin, 2021</xref>). Finally, the complete circular chloroplast genome map was visualized using OrganellarGenomeDRAW (OGDRAW) through the MPI-MP Chlorobox web server (<xref ref-type="bibr" rid="B10">Greiner et&#xa0;al., 2019</xref>), enabling clear graphical representation of gene content, orientation, and overall genome architecture.</p>
</sec>
</sec>
<sec id="s3">
<label>3</label>
<title>Data</title>
<sec id="s3_1">
<label>3.1</label>
<title>Characterization of <italic>Elaeodendron glaucum</italic> chloroplast genome</title>
<p>The complete chloroplast genome of <italic>E. glaucum</italic> was assembled as a circular molecule of 157,958 base pairs (bp) with an overall GC content of 37%. Its organization follows the typical quadripartite structure of angiosperm plastomes, consisting of a large single-copy (LSC) region of 86,485 bp with 35.21% GC content, a small single-copy (SSC) region of 18,363 bp with 31.80% GC content, and two inverted repeat (IR) regions of 26,555 bp each with 42.79% GC content (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>). The total genome size of <italic>E. glaucum</italic> is slightly larger than those reported for closely related genera within the Celastraceae family, including <italic>Celastrus vaniotii</italic> (H.L&#xe9;v.) Rehder (157,194 bp; GenBank accession: OR726632), <italic>E. kiautschovicus</italic> (157,611 bp; GenBank accession: PQ397793), <italic>Microtropis osmanthoides</italic> (Hand.-Mazz.) Hand.-Mazz. (156,659 bp; GenBank accession: NC 065714), and <italic>Parnassia faberi</italic> Oliv. (153,846 bp; GenBank accession: NC 061028), but shorter than <italic>Salacia menglaensis</italic> J.Y.Shen, L.C.Yan &amp; Landrein (163,255 bp; GenBank accession: NC 047214).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Circular gene map of the <italic>Elaeodendron glaucum</italic> chloroplast genome. Genes positioned on the inner track of the circular map are transcribed in the counterclockwise direction, while those on the outer track are transcribed clockwise. Distinct colors are used to indicate different functional categories of genes. The innermost circle illustrates the GC content in grey, with the lighter grey areas representing AT content.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1649473-g001.tif">
<alt-text content-type="machine-generated">Circular diagram of the Elaeodendron glaucum  chloroplast genome, totaling 157,958 base pairs. It features color-coded gene regions including photosystems, ATP synthase, and ribosomal proteins. Key regions like LSC, SSC, and IRB are marked. A legend at the bottom left explains the color codes for various gene types, such as photosystems in green and ribosomal RNAs in red.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Gene annotation of <italic>Elaeodendron glaucum</italic> chloroplast genome</title>
<p>A total of 133 genes were annotated within the <italic>E. glaucum</italic> chloroplast genome, encompassing 112 unique genes. These include 88 protein-coding genes (79 unique), 37 transfer RNA (<italic>t</italic>RNA) genes (29 unique), and 8 ribosomal RNA (<italic>r</italic>RNA) genes (4 unique). Among these, 17 genes contain a single intron, while three genes&#x2014;<italic>rps12</italic>, <italic>pafI</italic>, and <italic>clpP</italic>&#x2014;each possess two introns (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>).</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>List of genes in the <italic>Elaeodendron glaucum</italic> chloroplast genome.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Functional category</th>
<th valign="middle" align="left">Group of Gene</th>
<th valign="middle" align="left">Name of Gene</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" rowspan="7" align="left">Self-replication</td>
<td valign="middle" align="left">rRNA</td>
<td valign="middle" align="left">
<italic>rrn16<sup>d</sup>, rrn23<sup>d</sup>, rrn4.5<sup>d</sup>, rrn5<sup>d</sup>
</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">tRNA</td>
<td valign="middle" align="left">
<italic>trnH-GUG, trnK-UUU<sup>*</sup>, trnQ-UUG, trnS-GCU, trnG-GCC<sup>d*</sup>, trnR-UCU, trnC-GCA, trnD-GUC, trnY-GUA, trnE-UUC, trnT-GGU, trnS-UGA, trnfM-CAU<sup>d</sup>, trnS-GGA, trnT-UGU, trnL-UAA<sup>*</sup>, trnF-GAA, trnV-UAC<sup>*</sup>, trnW-CCA, trnP-UGG, trnI-CAU<sup>d</sup>, trnL-CAA<sup>d</sup>, trnV-GAC<sup>d</sup>, trnI-GAU<sup>d*</sup>, trnA-UGC<sup>d*</sup>, trnR-ACG<sup>d</sup>, trnN-GUU<sup>d</sup>, trnL-UAG</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">Large subunit ribosomal proteins (LSU)</td>
<td valign="middle" align="left">
<italic>rpl14, rpl16, rpl2<sup>d*</sup>, rpl20, rpl22, rpl23<sup>d</sup>, rpl32, rpl33, rpl36</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">Small subunit ribosomal proteins (SSU)</td>
<td valign="middle" align="left">
<italic>rps11, rps12<sup>d**</sup>, rps14, rps15, rps16, rps18, rps19 <sup>d</sup>, rps2, rps3, rps4, rps7 <sup>d</sup>, rps8</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">DNA dependent RNA polymerase</td>
<td valign="middle" align="left">
<italic>rpoA, rpoB, rpoC1<sup>*</sup>, rpoC2</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">Subunits of ATP synthase</td>
<td valign="middle" align="left">
<italic>atpA, atpB, atpE, atpF<sup>*</sup>, atpH, atpI</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">Subunits of NADH-dehydrogenase</td>
<td valign="middle" align="left">
<italic>ndhA<sup>*</sup>, ndhB<sup>d</sup>*, ndhC, ndhD, ndhE, ndhF, ndhG, ndhH, ndhI, ndhJ, ndhK</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">Photosynthesis</td>
<td valign="middle" align="left">Subunits of photosystem I</td>
<td valign="middle" align="left">
<italic>psaA, psaB, psaC, psaI, psaJ</italic>
</td>
</tr>
<tr>
<td valign="middle" rowspan="5" align="left">
</td>
<td valign="middle" align="left">Subunits of photosystem II</td>
<td valign="middle" align="left">
<italic>psbA, psbB, psbC, psbD, psbE, psbF, pdbH, psbI, psbJ, psbK, psbM, psbN, psbT, psbZ, pafI<sup>**</sup>
</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">Subunits of cytochrome b/f complex</td>
<td valign="middle" align="left">
<italic>petA, petB*, petD*, petG, petL, petN</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">Subunit rubisco</td>
<td valign="middle" align="left">
<italic>rbcL</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">Subunit of acetyl-CoA-carboxylase</td>
<td valign="middle" align="left">
<italic>accD</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">C-type cytochrome synthesis gene</td>
<td valign="middle" align="left">
<italic>ccsA</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">Other function</td>
<td valign="middle" align="left">Protease</td>
<td valign="middle" align="left">
<italic>clpP<sup>**</sup>
</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="left">Maturase</td>
<td valign="middle" align="left">
<italic>matK</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="left">Envelope membrane protein<break/>Translational initiation factor</td>
<td valign="middle" align="left">
<italic>cemA</italic>
<break/>
<italic>infA</italic>
</td>
</tr>
<tr>
<td valign="middle" align="left">Unknown function</td>
<td valign="middle" align="left">Conserved open reading frames</td>
<td valign="middle" align="left">
<italic>ycf1<sup>*</sup>, ycf15<sup>d</sup>, ycf2<sup>d</sup>, pafII</italic>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>d, gene duplication; *, single intron; **, double intron.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
</sec>
</body>
<back>
<sec id="s4" sec-type="data-availability">
<title>Data availability statement</title>
<p>The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found below: NCBI GenBank, accession PV153503.</p>
</sec>
<sec id="s5" sec-type="author-contributions">
<title>Author contributions</title>
<p>DSR: Writing &#x2013; review &amp; editing. IPAH: Writing &#x2013; review &amp; editing, Data curation. MM: Data curation, Writing &#x2013; review &amp; editing. IM: Formal analysis, Writing &#x2013; review &amp; editing, Methodology, Data curation. TYIW: Writing &#x2013; review &amp; editing. JTH: Writing &#x2013; review &amp; editing. DG: Writing &#x2013; review &amp; editing. NU: Writing &#x2013; review &amp; editing. R: Writing &#x2013; review &amp; editing. ASDI: Data curation, Writing &#x2013; review &amp; editing. RRI: Writing &#x2013; review &amp; editing. AS: Writing &#x2013; review &amp; editing. MRH: Data curation, Methodology, Conceptualization, Funding acquisition, Writing &#x2013; review &amp; editing, Formal analysis, Writing &#x2013; original draft.</p>
</sec>
<sec id="s6" 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 through the Joint Collaboration Rumah Program of the Organization for Research in Life and Environmental Sciences (ORHL), National Research and Innovation Agency (BRIN) for the 2023 fiscal year under contract number B-651/III.5/PR.03.06/2/2023.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>The authors express gratitude to the Directorate of Scientific Collection Management, as well as the management of Bogor Botanic Gardens, for their essential support and facilitation of this research.</p>
</ack>
<sec id="s7" 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="s8" 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>
<p>Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.</p>
</sec>
<sec id="s9" 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>
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