<?xml version="1.0" encoding="UTF-8" standalone="no"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing DTD v2.3 20070202//EN" "journalpublishing.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" article-type="research-article" dtd-version="2.3" xml:lang="EN">
<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.2025.1619267</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>Complete plastome and mitogenome assembly of endangered tree <italic>Karpatiosorbus bristoliensis</italic> reveals phylogenetic architecture for <italic>Sorbus</italic> sensu lato (Rosaceae)</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Li</surname>
<given-names>Qiang</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2905522/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/investigation/"/>
<role content-type="https://credit.niso.org/contributor-roles/supervision/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/resources/"/>
<role content-type="https://credit.niso.org/contributor-roles/software/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/project-administration/"/>
<role content-type="https://credit.niso.org/contributor-roles/validation/"/>
<role content-type="https://credit.niso.org/contributor-roles/formal-analysis/"/>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/visualization/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wei</surname>
<given-names>Ran</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/software/"/>
<role content-type="https://credit.niso.org/contributor-roles/visualization/"/>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>College of Life Sciences, Nanjing Forestry University</institution>, <addr-line>Nanjing, Jiangsu</addr-line>,&#xa0;<country>China</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>College of Manufacturing Engineering, Maanshan University</institution>, <addr-line>Maanshan</addr-line>,&#xa0;<country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Zhiqiang Wu, Chinese Academy of Agricultural Sciences, China</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Zhechen Qi, Zhejiang Sci-Tech University, China</p>
<p>Guilong Lu, Henan Institute of Science and Technology, China</p>
<p>Luan Rabelo, Vale Technological Institute (ITV), Brazil</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Qiang Li, <email xlink:href="mailto:liqiang9503@njfu.edu.cn">liqiang9503@njfu.edu.cn</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>08</day>
<month>07</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>16</volume>
<elocation-id>1619267</elocation-id>
<history>
<date date-type="received">
<day>27</day>
<month>04</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>09</day>
<month>06</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Li and Wei</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Li and Wei</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 genus <italic>Sorbus</italic> sensu lato (<italic>Sorbus</italic> s.l.) comprises over 260 species widely distributed across temperate regions of Asia, Europe, and North America. However, hybridization and polyploidization have posed significant challenges to phylogenetic and taxonomic studies within this genus.</p>
</sec>
<sec>
<title>Methods</title>
<p>Here, we assemble the first complete chloroplast and mitochondrial genomes of <italic>Karpatiosorbus bristoliensis</italic> to characterize organellar genomic features and establish a maternally inherited phylogenetic framework for <italic>Sorbus</italic> s.l.</p>
</sec>
<sec>
<title>Results and discussion</title>
<p>The mitochondrial genome of <italic>K. bristoliensis</italic> is circular (386,757 bp), encoding 55 genes, including 34 protein-coding genes, 18 rRNAs, and 3 tRNAs. Its chloroplast genome has a typical quadripartite structure (160,322 bp), containing 75 protein-coding genes, 29 tRNA genes, 4 rRNA genes, and one pseudogene (<italic>ycf1&#x3a8;</italic>). Homologous gene transfer analysis of <italic>Sorbus</italic> s.l. species revealed inter-organellar gene transfer ranging from 3,021 to 3,846 bp. RNA editing analysis revealed 274&#x2013;352 editing sites in <italic>Sorbus</italic> s.l., with <italic>nad4</italic> containing the greatest number of editing sites across all protein-coding genes except those in <italic>K. bristoliensis</italic>. Simple sequence repeat (SSR) analysis detected 48&#x2013;52 SSRs per species, predominantly mononucleotide repeats. Phylogenetic reconstruction on the basis of organellar genomes revealed that <italic>Karpatiosorbus</italic> is a sister to <italic>Torminalis</italic>. Plastome-based phylogeny revealed the non-monophyletic status of <italic>Sorbus</italic> s.s., attributed to the nested placement of the hybrid-origin genera <italic>Hedlundia</italic> and <italic>Scandosorbus</italic> within the genus. Additionally, <italic>Hedlundia austriaca</italic> and <italic>H. persica</italic> should be transferred to the nothogenus <italic>Sorbomeles</italic>. Mitochondrial genome collinearity analysis revealed extensive genomic structural rearrangements. Our findings not only delineate the structural characteristics of mitochondrial genomes across <italic>Sorbus</italic> s.l. taxa but also establish a high-resolution maternal phylogenetic framework for this genus.</p>
</sec>
</abstract>
<kwd-group>
<kwd>
<italic>Karpatiosorbus bristoliensis</italic>
</kwd>
<kwd>
<italic>Sorbus</italic> s.l.</kwd>
<kwd>mitochondrial genome</kwd>
<kwd>RNA editing events</kwd>
<kwd>gene transfer</kwd>
<kwd>phylogenetic analysis</kwd>
</kwd-group>
<counts>
<fig-count count="7"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="77"/>
<page-count count="14"/>
<word-count count="5977"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Plant Bioinformatics</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<title>Introduction</title>
<p>The genus <italic>Sorbus</italic> L. were primarily established by Linnaeus and included only two species (<italic>S. aucuparia</italic> L. and <italic>S. domestica</italic> L.) (<xref ref-type="bibr" rid="B26">Linnaeus, 1753</xref>). Subsequently, more species were transferred to this genus. <italic>Sorbus</italic> L. broadly (<italic>Sorbus</italic> L. sensu lato) contains approximately 260 species characterized by both simple-leaved species and pinnately compound-leaved species (<xref ref-type="bibr" rid="B70">Y&#xfc; and Lu, 1974</xref>; <xref ref-type="bibr" rid="B43">Phipps et&#xa0;al., 1990</xref>; <xref ref-type="bibr" rid="B31">Lu and Stephen, 2003</xref>). <italic>Sorbus</italic> s.l. comprises trees or shrubs typically restricted to high altitudes, occupying diverse habitats including mountain valleys, streamsides, rocky slopes, and scrublands (<xref ref-type="bibr" rid="B31">Lu and Stephen, 2003</xref>). These plants are widely distributed across temperate Asia, Europe, and North America (<xref ref-type="bibr" rid="B43">Phipps et&#xa0;al., 1990</xref>). Many species exhibit ornamental value due to their conspicuous colorful fruits (<xref ref-type="bibr" rid="B35">McAllister, 2005</xref>).</p>
<p>Previous molecular and morphological evidence suggests that <italic>Sorbus</italic> s.l. is paraphyletic and raises six subgenera to the generic rank (<italic>Aria</italic> (Pers.) Host, <italic>Cormus</italic> Spach<italic>, Chamaemespilus</italic> Medikus<italic>, Micromeles</italic> Decaisne, <italic>Sorbus</italic> sensu stricto (<italic>Sorbus</italic> s.s.), and <italic>Torminalis</italic> Medikus) (<xref ref-type="bibr" rid="B5">Campbell et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B76">Zheng and Zhang, 2007</xref>; <xref ref-type="bibr" rid="B44">Potter et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B30">Lo and Donoghue, 2012</xref>). Among the six genera, the genera <italic>Aria</italic>, <italic>Chamaemespilus</italic>, <italic>Micromeles</italic>, and <italic>Torminalis</italic> all have simple leaves, whereas <italic>Cormus</italic> and <italic>Sorbus</italic> s.s. are characterized by compound leaves. However, the generic boundaries of these taxa have been continuously revised. For instance, <xref ref-type="bibr" rid="B36">Mezhenska et&#xa0;al. (2018)</xref> proposed transferring all Asian simple-leaved species to the genus <italic>Micromeles</italic>, including the previously taxonomically disputed genera <italic>Aria</italic> and <italic>Micromeles</italic>. In the same year, <xref ref-type="bibr" rid="B51">Rushforth (2018)</xref> alternatively classified Asian simple-leaved species within <italic>Micromeles</italic> and five newly established genera (<italic>Griffitharia, Alniaria, Thomsonaria, Dunnaria, and Wilsonaria</italic>) on the basis of morphological characteristics.</p>
<p>The primary factors contributing to the taxonomic complexity within <italic>Sorbus</italic> s.l. are hybridization and polyploidization. Notably, previous works highlighted that polyploidization, frequent natural hybridization and apomixis play crucial roles in the ongoing genetic diversification of <italic>Sorbus</italic> s.l. species (<xref ref-type="bibr" rid="B49">Robertson et&#xa0;al., 2010</xref>). Biparental, triparental and multiple-hybrid origins (<xref ref-type="bibr" rid="B40">N&#xe9;meth et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B39">Nelson-Jones et&#xa0;al., 2002</xref>; <xref ref-type="bibr" rid="B48">Robertson et&#xa0;al., 2004</xref>) contribute to allodiploid, triploid and tetraploid species, rendering the taxonomy complex. In addition, interspecific hybridization also promotes the innovation of morphological characteristics, such as the number of pairs of leaflets and fruit color (<xref ref-type="bibr" rid="B39">Nelson-Jones et&#xa0;al., 2002</xref>; <xref ref-type="bibr" rid="B66">Wu et&#xa0;al., 2024</xref>).</p>
<p>The taxonomic treatment of numerous hybrid-origin species remains unresolved, particularly with respect to their delimitation and placement within phylogenetic frameworks. These hybrid-origin species have been taxonomically elevated to subgenera (<xref ref-type="bibr" rid="B46">Rich et&#xa0;al., 2014</xref>) or genera (<xref ref-type="bibr" rid="B52">Sennikov and Kurtto, 2017</xref>). For example, <xref ref-type="bibr" rid="B52">Sennikov and Kurtto (2017)</xref> elevated certain hybrid-origin taxa within the genus <italic>Sorbus</italic> to five hybridogenous genera (<italic>Borkhausenia</italic> = <italic>Aria</italic> &#xd7; <italic>Sorbus</italic> &#xd7; <italic>Torminalis</italic>, <italic>Hedlundia</italic> = <italic>Aria</italic> &#xd7; <italic>Sorbus</italic>, <italic>Karpatiosorbus</italic> = <italic>Aria</italic> &#xd7; <italic>Torminalis</italic>, <italic>Majovskya</italic> = <italic>Aria</italic> &#xd7; <italic>Chamaemespilus</italic>, <italic>Normeyera</italic> = <italic>Aria</italic> &#xd7; <italic>Chamaemespilus</italic> &#xd7; <italic>Sorbus</italic>).</p>
<p>
<italic>Karpatiosorbus</italic> has a hybrid origin from <italic>Aria</italic> (Pers.) Host &#xd7; <italic>Torminalis</italic> Medik, including one sexual hybrid (<italic>Karpatiosorbus</italic> &#xd7; <italic>hybrida</italic> (Borkh.) Sennikov &amp; Kurtto) and 84 apomictic species (<xref ref-type="bibr" rid="B52">Sennikov and Kurtto, 2017</xref>). Among these 85 species, <italic>Karpatiosorbus bristoliensis</italic> (Wilmott) Sennikov &amp; Kurtto is a shrub or small tree approximately 30 ft high and has restricted areas of distribution in Avon Gorge, England. It inhabits open limestone outcrops, slopes, scrub, open grassland, quarry edges, and ancient woodland plateau soils, primarily within the Leigh Woods ecosystem (<xref ref-type="bibr" rid="B47">Rich et&#xa0;al., 2022</xref>). Despite its ornamental value, this species has been assessed as &#x2018;Endangered&#x2019; under criterion D (<xref ref-type="bibr" rid="B18">IUCN, 2001</xref>) and on the Global Red List owing to its small population size (<xref ref-type="bibr" rid="B47">Rich et&#xa0;al., 2022</xref>). Using six chloroplast fragments, <xref ref-type="bibr" rid="B6">Chester et&#xa0;al. (2007)</xref> revealed that <italic>K. bristoliensis</italic> and <italic>Sorbus torminalis</italic> (L.) Crantz shared an identical haplotype (haplotype B), while proposing that <italic>K. bristoliensis</italic> may represent a case of <italic>in situ</italic> speciation. Employing microsatellite analysis, <xref ref-type="bibr" rid="B49">Robertson et&#xa0;al. (2010)</xref> determined the triploid hybrid origin of <italic>K. bristoliensis</italic> with diploid <italic>S. torminalis</italic> as the maternal progenitor and tetraploid <italic>S. eminens</italic> E.F.Warb. as the paternal progenitor (<xref ref-type="bibr" rid="B49">Robertson et&#xa0;al., 2010</xref>). Previous studies utilizing organellar genome have provided novel insights into the genetic diversity and conservation of endangered species (<xref ref-type="bibr" rid="B74">Zhang et&#xa0;al., 2024</xref>). However, the limited genomic information available for this endangered species hinders our understanding of its genomic structure and genomic diversity. The organellar genomes assembled in this study will lay the foundation for conservation genomics in <italic>K. bristoliensis</italic>.</p>
<p>The mitochondrial genome has received increasing attention in the field of plant phylogenomics (<xref ref-type="bibr" rid="B25">Lin et&#xa0;al., 2025</xref>). This is because of the slow evolutionary rate of mitochondrial genes compared with nuclear genes. Unlike the chloroplast genome, plant mitochondrial genomes are large (66 kb to 18.99 Mb) (<xref ref-type="bibr" rid="B55">Skippington et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B15">Huang et&#xa0;al., 2024a</xref>), exhibit frequent recombination, and often incorporate foreign DNA via horizontal gene transfer (HGT) from chloroplasts and nuclear (<xref ref-type="bibr" rid="B58">Wang et al., 2025</xref>; <xref ref-type="bibr" rid="B75">Zhang et&#xa0;al., 2025</xref>). These features result in highly variable genome sizes and architectures across plant lineages (<xref ref-type="bibr" rid="B12">Gualberto and Newton, 2017</xref>; <xref ref-type="bibr" rid="B64">Wu et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B33">Lu et&#xa0;al., 2025</xref>). The plant mitochondrial genome harbors abundant repetitive sequences, including simple sequence repeats (SSRs), tandem repeats, and interspersed repeats. These repetitive elements serve as a foundation for the development of diverse molecular markers to investigate population genetic diversity (<xref ref-type="bibr" rid="B38">Morgante et&#xa0;al., 2002</xref>). Moreover, the high abundance of repeats contributes significantly to the complex structural rearrangements observed in mitochondrial genomes, such as recombination-mediated inversions, duplications, and the formation of multipartite architectures (<xref ref-type="bibr" rid="B3">Bi et&#xa0;al., 2022</xref>).</p>
<p>The availability of extensive chloroplast genome sequencing data for the genus <italic>Sorbus</italic> s.l. enables the utilization of these datasets to address unresolved phylogenetic questions within this taxonomic group. Furthermore, recent advances in long-read sequencing have resolved complex plant mitochondrial genome structures. Taking advantage on the previously released PacBio HiFi sequencing data of <italic>Karpatiosorbus bristoliensis</italic> in the NCBI database, we utilized these datasets to perform the first <italic>de novo</italic> assembly of both chloroplast and mitochondrial genomes for the hybrid genus <italic>Karpatiosorbus</italic>. Our main goals are to (1) assemble and characterize the first mitochondrial and plastid genomes of this species; (2) compare the mitochondrial genomes of <italic>Sorbus</italic> s.l. species; (3) explore repetitive sequences and RNA editing sites; and (4) reconstruct the maternal phylogenetic framework within <italic>Sorbus</italic> s.l. utilizing organellar genomes.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<title>Materials and methods</title>
<sec id="s2_1">
<title>Plant material, mitochondrial genome assembly and annotation</title>
<p>We obtained PacBio-HiFi data for <italic>Karpatiosorbus bristoliensis</italic> (ERR13245294) from the SRA (<ext-link ext-link-type="uri" xlink:href="https://www.ncbi.nlm.nih.gov/sra/?term=ERR13245294">https://www.ncbi.nlm.nih.gov/sra/?term=ERR13245294</ext-link>). The whole-genome sequencing of <italic>K. bristoliensis</italic> revealed a total of 50.3G bases. For mitogenome assembly, we employed PMAT v2.1.2 (<ext-link ext-link-type="uri" xlink:href="https://github.com/aiPGAB/PMAT2/releases">https://github.com/aiPGAB/PMAT2/releases</ext-link>) (<xref ref-type="bibr" rid="B4">Bi et&#xa0;al., 2024</xref>), an efficient plant mitogenome assembly toolkit using low-coverage HiFi sequencing data, to perform <italic>de novo</italic> assembly of the <italic>K. bristoliensis</italic> mitogenome with the command &#x201c;PMAT autoMito -I -o -st -g -m&#x201d;. For plastid genome assembly, the ptGAUL v1.0.5 pipeline (<ext-link ext-link-type="uri" xlink:href="https://github.com/Bean061/ptgaul">https://github.com/Bean061/ptgaul</ext-link>) (<xref ref-type="bibr" rid="B77">Zhou et&#xa0;al., 2023</xref>) was applied to extract and assemble the plastid genome from PacBio-HiFi long-read data with standard parameters. We employed Bandage (<xref ref-type="bibr" rid="B63">Wick et&#xa0;al., 2015</xref>) to visualize the graphical representation of the mitochondrial and plastid genomes produced by the PMAT and ptGAUL pipelines, respectively. Mitogenome annotation was performed via the online program Plant Mitochondrial Genome Annotator (PMGA) (<ext-link ext-link-type="uri" xlink:href="http://47.96.249.172:16084/index.html">http://47.96.249.172:16084/index.html</ext-link>), with the database set as 319 mitogenomes (<xref ref-type="bibr" rid="B22">Li et&#xa0;al., 2024</xref>). This database uses multiple sequence alignment to identify genes. The plastid genome annotation was performed via the online program CPGAVAS2 (<ext-link ext-link-type="uri" xlink:href="http://47.96.249.172:16019/analyzer/home">http://47.96.249.172:16019/analyzer/home</ext-link>) (<xref ref-type="bibr" rid="B53">Shi et&#xa0;al., 2019</xref>), with <italic>Sorbus helenae</italic> (NC_068536.1) as the reference genome. Manual corrections were performed via Sequin software. We utilized OGDRAW (v.1.3.1) (<ext-link ext-link-type="uri" xlink:href="https://chlorobox.mpimp-golm.mpg.de/ogdraw.html">https://chlorobox.mpimp-golm.mpg.de/ogdraw.html</ext-link>) to generate plastid genome and mitogenome maps.</p>
</sec>
<sec id="s2_2">
<title>Analysis of repeat sequences</title>
<p>The prediction of simple sequence repeats (SSRs) was performed using the online program MISA-web (<ext-link ext-link-type="uri" xlink:href="https://www.web-blast.ipk-gatersleben.de/misa/">https://www.web-blast.ipk-gatersleben.de/misa/</ext-link>). The parameters of mono-, di-, tri-, tetra-, penta-, and hexanucleotides were set to 10, 5, 4, 3, 3, and 3, respectively. Tandem repeats within the five species were detected using Tandem Repeats Finder v4.09 (<xref ref-type="bibr" rid="B2">Benson, 1999</xref>) with default parameters. Dispersed repeats were identified using the online REPuter program (<ext-link ext-link-type="uri" xlink:href="https://bibiserv.cebitec.uni-bielefeld.de/reputer">https://bibiserv.cebitec.uni-bielefeld.de/reputer</ext-link>) (<xref ref-type="bibr" rid="B21">Kurtz, 2001</xref>) with a minimal repeat size of 30 bp and a Hamming distance of 3.</p>
</sec>
<sec id="s2_3">
<title>Homologous fragment analysis</title>
<p>we employed the online BLAST tool (<ext-link ext-link-type="uri" xlink:href="https://blast.ncbi.nlm.nih.gov/Blast.cgi">https://blast.ncbi.nlm.nih.gov/Blast.cgi</ext-link>) to search highly similar sequences between the mitogenome and plastid genomes of <italic>K. bristoliensis, M. alnifolia</italic> (Siebold &amp; Zucc.) Koehne, T. <italic>glaberrima</italic> (Gand.) Sennikov &amp; Kurtto<italic>, S. aucuparia</italic> L., and <italic>Sorbus aucuparia</italic> subsp. <italic>pohuashanensis</italic> (Hance) McAll., respectively. The identification parameters for homologous sequences were set to &#x2265;70% minimum similarity and &#x2264;1e-5 E-value threshold.</p>
</sec>
<sec id="s2_4">
<title>Detection of RNA editing events within <italic>Sorbus</italic> s.l.</title>
<p>We employed the online program PMGA (<ext-link ext-link-type="uri" xlink:href="http://47.96.249.172:16084/deepredmt.html">http://47.96.249.172:16084/deepredmt.html</ext-link>) to predict RNA editing events in the mitogenomes of <italic>K. bristoliensis, M. alnifolia, T. glaberrima, S. aucuparia, and S. aucuparia subsp. pohuashanensis</italic>. This online program uses Deepred-Mt (<xref ref-type="bibr" rid="B8">Edera et&#xa0;al., 2021</xref>), a novel deep convolutional neural network, to detect potential RNA editing events. The cut-off value was set to 0.5 for accurate prediction.</p>
</sec>
<sec id="s2_5">
<title>Nucleotide diversity analysis</title>
<p>For plastid genome analyses, plastid genome sequences from a total of 59 species of <italic>Sorbus</italic> s.l. were downloaded. We used the command &#x201c;cpstools Pi -d work_dir&#x201d; from the CPStools package (<xref ref-type="bibr" rid="B16">Huang et&#xa0;al., 2024b</xref>) to calculate Pi values for the analyzed species. This package automatically extracts shared gene regions and intergenic sequences from GenBank-format files in work_dir, performs multiple-sequence alignments, and computes Pi values. For mitogenome analyses, we utilized PhyloSuite v1.2.3 (<xref ref-type="bibr" rid="B72">Zhang et&#xa0;al., 2020</xref>) to extract protein-coding genes from the five <italic>Sorbus</italic> s.l. species using default parameters. A total of 35 shared protein-coding genes were aligned using MAFFT (<ext-link ext-link-type="uri" xlink:href="https://mafft.cbrc.jp/alignment/server/">https://mafft.cbrc.jp/alignment/server/</ext-link>) (<xref ref-type="bibr" rid="B20">Katoh et&#xa0;al., 2018</xref>) with the default parameters. The nucleotide diversity (Pi) of each gene was subsequently calculated using DnaSP v6 (<xref ref-type="bibr" rid="B50">Rozas et&#xa0;al., 2017</xref>) with the option &#x201c;compute variance of Pi&#x201d;.</p>
</sec>
<sec id="s2_6">
<title>Phylogenetic analysis</title>
<p>The mitochondrial phylogenomic tree was reconstructed via 14 mitochondrial genomes, including those of five <italic>Sorbus</italic> s.l. species (<italic>Sorbus aucuparia</italic> subsp. <italic>pohuashanensis</italic> ON478177, <italic>Sorbus aucuparia</italic> MT648825, <italic>Micromeles alnifolia</italic> PP506330, <italic>Torminalis glaberrima</italic> MT610102, and <italic>Karpatiosorbus bristoliensis</italic>), with <italic>Rosa rugosa</italic> (PQ474155) designated as the outgroup. We employed PhyloSuite v1.2.3 (<xref ref-type="bibr" rid="B72">Zhang et&#xa0;al., 2020</xref>) to extract protein-coding genes that were shared by the 14 species. Multiple sequence alignment was subsequently performed using MAFFT (<ext-link ext-link-type="uri" xlink:href="https://mafft.cbrc.jp/alignment/server/">https://mafft.cbrc.jp/alignment/server/</ext-link>) (<xref ref-type="bibr" rid="B20">Katoh et&#xa0;al., 2018</xref>) with default parameters. The aligned sequence trimming was performed using the trimAl program within the PhyloSuite v1.2.3 (<xref ref-type="bibr" rid="B72">Zhang et&#xa0;al., 2020</xref>) with the default (-automated1) parameter. We performed phylogenetic reconstruction using maximum likelihood method in IQ-TREE v2 (<xref ref-type="bibr" rid="B37">Minh et&#xa0;al., 2020</xref>), with 1,000 bootstrap replicates (-bb 1000) and 1,000 replicates for the SH-like approximate likelihood ratio test (SH-aLRT; -alrt 1000). The optimal nucleotide substitution model was determined using ModelFinder (<xref ref-type="bibr" rid="B19">Kalyaanamoorthy et&#xa0;al., 2017</xref>) in IQ-TREE v2, which identified K3Pu+F+R4 as the best-fit model based on the Bayesian Information Criterion (BIC).</p>
<p>A total of 65 plastomes, including those of 59 <italic>Sorbus</italic> s.l. species, were utilized for phylogenetic analyses (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S1</bold>
</xref>). The whole plastid genomes were aligned with MAFFT (<ext-link ext-link-type="uri" xlink:href="https://mafft.cbrc.jp/alignment/server/">https://mafft.cbrc.jp/alignment/server/</ext-link>) (<xref ref-type="bibr" rid="B20">Katoh et&#xa0;al., 2018</xref>) with default parameter. Phylogenetic reconstruction employed identical software and parameters as above, with the TIM+F+R4 nucleotide substitution model specified according to the Bayesian Information Criterion (BIC)</p>
</sec>
<sec id="s2_7">
<title>Collinearity analysis</title>
<p>We performed pairwise sequence alignments among five species of the <italic>Sorbus</italic> s.l genus using MUMmer v4.0.1 (<xref ref-type="bibr" rid="B34">Mar&#xe7;ais et&#xa0;al., 2018</xref>), with the parameters set as: nucmer &#x2013;maxmatch -c 100 -b 500 -l 50. Filtering of aligned sequences was performed with the following parameters: a minimum alignment identity of 90% and a minimum alignment length of 50 bp. Identification of collinear sequences and structural rearrangement events among mitochondrial genomes was conducted using SyRI v1.7.0 (<xref ref-type="bibr" rid="B11">Goel et&#xa0;al., 2019</xref>). Genome structural visualization was generated using Plotsr v1.1.0 (<xref ref-type="bibr" rid="B10">Goel and Schneeberger, 2022</xref>).</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<title>Results</title>
<sec id="s3_1">
<title>General features of <italic>Karpatiosorbus bristoliensis</italic> mitogenome and plastome</title>
<p>We employed the autoMito program in PMAT to assemble the mitochondrial genome of <italic>K. bristoliensis</italic>. The average depth of the assembled results was 1408.7x (<xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Figure S1</bold>
</xref>). The mitochondrial genome of <italic>K. bristoliensis</italic> was 386,757 bp in size. It encoded 55 unique genes, including 34 protein-coding genes, 18 tRNA genes and 3 rRNA genes (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1A</bold>
</xref>). The GC content was 45.3%. Three genes (<italic>ccmFC, trnE-UUC, trnS-GCU</italic>) had one intron, one gene (<italic>nad4</italic>) had three introns, and four genes (<italic>nad1, nad2, nad5, nad7</italic>) had four introns (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Tables S2</bold>
</xref>, <xref ref-type="supplementary-material" rid="SM1">
<bold>S3</bold>
</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Mitochondrial and chloroplast genome diagram of <italic>K</italic>. <italic>bristoliensis</italic>. <bold>(A)</bold> mitogenome; <bold>(B)</bold> Plastome. Genes encoded on the forward strand are positioned on the outer circumference, while those on the reverse strand are located on the inner circumference. The gray inner ring represents the GC content.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1619267-g001.tif">
<alt-text content-type="machine-generated">Circular diagrams depict the mitochondrial and chloroplast genomes of Karpatiosorbus bristoliensis. Diagram A shows the mitochondrial genome with 386,757 base pairs, highlighting various genes and their functions in different colors. Diagram B illustrates the chloroplast genome with 160,322 base pairs, also color-coded for gene identification. Both diagrams include labeled segments representing gene categories, with legends for function types, such as transfer RNAs and ribosomal RNAs.</alt-text>
</graphic>
</fig>
<p>The <italic>K. bristoliensis</italic> plastid genome has structures identical to those of most angiosperm plants, consisting of one large single copy (LSC), one small single copy (SSC), and two inverted repeats (IRs) (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1B</bold>
</xref>). The total length of the plastid genome was 160,322 bp, including 88,182 bp, 19,306 bp and 26,417 bp in the LSC, SSC and IR regions, respectively. The GC content was 36.50%. The plastid genome encodes 75 protein-coding genes, 29 tRNA genes, four rRNA genes, and one pseudogene (<italic>ycf1</italic>&#x3a8;) for a total of 108 unique genes. Among these genes, eight genes (<italic>rps16, atpF, rpoC1, petB, petD, ndhB, rpl2, ndhA</italic>) have one intron, and two genes (<italic>ycf3, clpP</italic>) possess two introns (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S4</bold>
</xref>).</p>
</sec>
<sec id="s3_2">
<title>Repeat sequence analyses</title>
<p>We detected 52, 53, 52, 48, and 51 simple sequence repeats (SSRs) in the mitochondrial genomes of <italic>K. bristoliensis, M. alnifolia, S. aucuparia</italic> subsp. <italic>pohuashanensis, S. aucuparia</italic>, and <italic>T. glaberrima</italic>, respectively (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2A</bold>
</xref>). All five species possess three types of SSRs: monomeric, dimeric, and trimeric repeats, with monomeric repeats being the most abundant. Notably, most SSRs were distributed in intergenic spacer regions, with fewer detected in protein-coding regions (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S5</bold>
</xref>). Within protein-coding regions, SSRs primarily occurred in <italic>nad1</italic>, <italic>nad2</italic>, and <italic>nad3</italic> genes (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2B</bold>
</xref>). We identified 50 dispersed repeats in each species, consisting of forward and palindromic repeats (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2C</bold>
</xref>). Repeat sequence lengths ranged from 91&#x2013;428 bp (<italic>K. bristoliensis</italic>), 88&#x2013;2745 bp (<italic>M. alnifolia</italic>), 76&#x2013;26,130 bp (<italic>S. aucuparia</italic> subsp. <italic>pohuashanensis</italic>), 86&#x2013;6452 bp (<italic>S. aucuparia</italic>), and 80&#x2013;438 bp (<italic>T. glaberrima</italic>) (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2C</bold>
</xref>). Notably, <italic>S. aucuparia</italic> contained two large palindromic repeats (26,130 bp and 24,730 bp; <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S6</bold>
</xref>). A total of 24, 15, 22, 22, and 24 tandem repeats were identified in the mitochondrial genomes of <italic>K. bristoliensis, M. alnifolia, S. aucuparia</italic> subsp<italic>. pohuashanensis, S. aucuparia</italic>, and <italic>T. glaberrima</italic>, respectively (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2D</bold>
</xref>) (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S7</bold>
</xref>).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Distribution of repeat sequence in <italic>Sorbus</italic> s.l. mitogenome. <bold>(A)</bold> the number of SSRs in mitogenome; <bold>(B)</bold> distribution of SSRs in mitogenome; <bold>(C)</bold> the number of dispersed repeats; <bold>(D)</bold> the number of tandem repeats.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1619267-g002.tif">
<alt-text content-type="machine-generated">Panel A shows a bar chart of the number of SSRs in different species, categorized as monomeric, dimeric, and trimeric. Panel B presents the number of SSRs in intergenetic spacers and protein-coding regions. Panel C displays forward and palindromic repeat numbers in various species. Panel D depicts the number of tandem repeats in different species.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s3_3">
<title>Analysis of inter-organellar gene transfer events</title>
<p>Mitochondrial genome analyses revealed 14, 13, 12, 12, and 14 chloroplast-derived DNA fragments in <italic>K. bristoliensis</italic>, <italic>M. alnifolia</italic>, <italic>T. glaberrima</italic>, <italic>S. aucuparia</italic>, and <italic>S. aucuparia</italic> subsp. <italic>pohuashanensis</italic>, respectively (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S8</bold>
</xref>). The lengths of these transferred sequences exhibited limited variation, ranging from 3,846 bp (<italic>K. bristoliensis</italic>), 3,834 bp (<italic>M. alnifolia</italic>), 3,830 bp (<italic>T. glaberrima</italic>), 3,755 bp (<italic>S. aucuparia</italic>), and 3,021 bp (<italic>S. aucuparia</italic> subsp. <italic>pohuashanensis</italic>). Notably, the largest individual transferred fragment spanned 1,874 bp. Notably, <italic>K. bristoliensis</italic>, <italic>T. glaberrima</italic>, and <italic>S. aucuparia</italic> subsp<italic>. pohuashanensis</italic> each contained five fully transferred tRNA genes (<italic>trnW-CCA</italic>, <italic>trnD-GUC</italic>, <italic>trnH-GUG</italic>, <italic>trnN-GUU</italic>, and <italic>trnM-CAU</italic>). In contrast, <italic>M. alnifolia</italic> and <italic>S. aucuparia</italic> exhibited four completely transferred tRNA genes (<italic>trnW-CCA</italic>, <italic>trnH-GUG</italic>, <italic>trnD-GUC</italic>, and <italic>trnN-GTT</italic>). Furthermore, partial transfer events involving protein-coding genes were detected, including fragments of <italic>psbE</italic>, <italic>psbC</italic>, <italic>psaA</italic>, <italic>psbA</italic>, and <italic>atpA.</italic>
</p>
</sec>
<sec id="s3_4">
<title>Prediction of RNA editing sites</title>
<p>A total of 442, 403, 474, 463, and 352 RNA editing sites were identified in the mitochondrial genomes of <italic>K. bristoliensis</italic>, <italic>M. alnifolia</italic>, <italic>S. aucuparia</italic>, <italic>S. aucuparia</italic> subsp. <italic>pohuashanensis</italic>, and <italic>T. glaberrima</italic>, respectively (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>). Comparative analysis revealed that three taxa (<italic>K. bristoliensis</italic>, <italic>S. aucuparia</italic>, and <italic>S. aucuparia</italic> subsp. <italic>pohuashanensis</italic>) each contained RNA editing modifications in 32 protein-coding genes, while <italic>M. alnifolia</italic> and <italic>T. glaberrima</italic> possessed 31 and 28 edited genes, respectively. Notably, <italic>nad4L</italic> exhibited the lowest frequency of RNA editing events among all protein-coding genes across the five species, each containing a single editing site. The <italic>nad4</italic> gene demonstrated the highest RNA editing activity in four mitochondrial genomes (<italic>M. alnifolia, S. aucuparia, S. aucuparia</italic> subsp. <italic>pohuashanensis</italic>, and <italic>T. glaberrima</italic>), with 39 editing sites identified in <italic>M. alnifolia</italic>. In contrast, the <italic>K. bristoliensis</italic> mitochondrial genome showed unique editing patterns, where <italic>ccmB</italic> and <italic>ccmC</italic> collectively contained the highest number of RNA editing sites (31 total), representing a distinct regulatory feature in this species. All RNA editing sites belong to C to U. Notably, the start codons of <italic>cox1, nad1, rpl16</italic>, and <italic>rps4</italic> in the mitochondrial genome <italic>of K. bristoliensis</italic>, through RNA editing, have changed from ACG to AUG. Additionally, RNA editing has contributed to stop codons for <italic>atp6</italic> in the mitochondrial genomes of <italic>K. bristoliensis</italic>, <italic>S. aucuparia</italic>, and <italic>S. aucuparia</italic> subsp<italic>. pohuashanensis</italic> (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S9</bold>
</xref>). The most frequent amino acid substitution was serine-to-leucine, except in <italic>M. alnifolia</italic> where proline-to-leucine predominated (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3B</bold>
</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>The number of RNA editing sites identified in the mitogenomes. <bold>(A)</bold> the number of RNA editing sites in <italic>Sorbus</italic> s.l. mitogenome; <bold>(B)</bold> number of types of RNA editing.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1619267-g003.tif">
<alt-text content-type="machine-generated">Two bar graphs show RNA editing in five species: *K. bristolensis*, *T. glaberrima*, *A. alnifolia*, *S. aucuparia*, and *S. aucuparia* subsp. *pohuashanensis*. Graph A displays RNA editing numbers per gene site, with *ndhB* showing the highest editing. Graph B details amino acid changes, with Pro to Ser and Ser to Phe being the most frequent. Each species is represented by a different colored bar.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s3_5">
<title>Nucleotide diversity analysis</title>
<p>Nucleotide diversity analysis was performed using DnaSP v6.0. For mitochondrial genomes, the Pi values ranged from 0 to 0.01808, with the <italic>nad4</italic> gene exhibiting the highest diversity (0.01808) (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4A</bold>
</xref>). In plastid genomes, the Pi values showed greater conservation, spanning from 0 to 0.02672 (<italic>petG-trnW-CCA</italic>) (<xref ref-type="fig" rid="f4">
<bold>Figures 4B&#x2013;D</bold>
</xref>). Comparative analysis identified seven hypervariable regions as potential molecular markers. Seven regions with relatively high Pi values may be designed for DNA barcodes, including <italic>trnH-GUG-psbA</italic>, <italic>trnT-GUG-trnL-UAA</italic>, <italic>petG-trnW-CCA</italic>, <italic>ndhC-trnV-UAC-exon2</italic>, <italic>trnW-CCA-trnP-UGG</italic>, <italic>rpl33-rps18</italic>.</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Nucleotide diversity analysis of <italic>Sorbus</italic> s.l. species. <bold>(A)</bold> mitogenome; <bold>(B&#x2013;D)</bold> plastome.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1619267-g004.tif">
<alt-text content-type="machine-generated">Line graphs labeled A, B, and C show Pi values for various gene sequences on the x-axes. Graph A has peaks at &#x201c;psbK&#x201d; and &#x201c;petL,&#x201d; B peaks at &#x201c;trnH-GUG-psbA,&#x201d; and C peaks at &#x201c;petG-trnW-CCA.&#x201d; The y-axes represent Pi value scales from 0 to 0.0045, 0.02, and 0.035 respectively.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s3_6">
<title>Phylogenomic analyses</title>
<p>We recovered a plastome-based phylogeny of <italic>Sorbus</italic> s.l., which included an ingroup of 59 accessions (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>). Our plastome-based phylogeny yielded strong bootstrap values (&gt; 90%) for the vast majority of branches. With robust node support, we recovered eight major clades (A, B, C, D, E, F, G, H). Among these clades, <italic>Sorbus</italic> s.l. was dispersed in six clades. Clade A contains four genera (<italic>Chamaemespilus</italic>, <italic>Aria</italic>, <italic>Torminalis</italic>, and <italic>Karpatiosorbus</italic>) of <italic>Sorbus</italic> s.l. <italic>Karpatiosorbus</italic>, which are closely phylogenetically related to <italic>Torminalis</italic>. The genus <italic>Aria</italic> was found to be nonmonophyletic, with one species embedded in the genus <italic>Chamaemespilus</italic>. Clade C contains one species, <italic>Cormus domestica.</italic> Clade E corresponds to the genera <italic>Micromeles</italic> and <italic>Hedlundia</italic>. <italic>Micromeles</italic> are not monophyletic, with <italic>Hedlundia</italic> embedded in them. Clades F-H correspond to <italic>Sorbus</italic> s.s. and are expected to include the hybrid-origin genus <italic>Hedlundia</italic> and <italic>Scandosorbus</italic>.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>The maximum likelihood phylogeny reconstructed with IQ-TREE v2 displays bootstrap values on branches. Colors in the figure denote <italic>Sorbus</italic> s.l. lineages, with green designating <italic>Sorbus</italic> s.s. clades. Botanical images beneath the phylogeny represent constituent genera of <italic>Sorbus</italic> s.l. included in this study, with parenthetical notation indicating documented hybrid origins.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1619267-g005.tif">
<alt-text content-type="machine-generated">Phylogenetic tree highlighting relationships within Sorbus species, depicted in distinct clades with statistical support values. Below, there are images of different Sorbus species and their hybrids, each bearing unique berries and leaves. Species names such as Hedlundia, Micromeles, and Karpatiosorbus are labeled alongside their respective plant images.</alt-text>
</graphic>
</fig>
<p>To better understand the mitochondrial genome evolution of <italic>Sorbus</italic> s.l., we also utilized the maximum likelihood method to construct mitochondrial genome-based phylogenetic trees based on 35 shared protein-coding genes (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>). The <italic>Prunus</italic> genus was used as the outgroup. Most nodes of the phylogenetic tree presented relatively lower bootstrap values than those of the plastome-based phylogenetic tree did. Consistent with the plastome-based phylogeny, the genus <italic>Karpatiosorbus</italic> is a sister to <italic>Torminalis</italic>.</p>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>ML phylogeny (IQ-TREE v2; 34 shared protein-coding genes) showing bootstrap values on branches.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1619267-g006.tif">
<alt-text content-type="machine-generated">Phylogenetic tree depicting relationships among various plant species. Includes Alniaria alnifolia, Crataegus bretschneideri, and others, with Karpatiosorbus bristoliensis highlighted. Bootstrap values indicate support for nodes, with Rosa rugosa as an outgroup.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s3_7">
<title>Analysis of mitogenome collinearity and structural rearrangement</title>
<p>Analysis of mitochondrial genome sequences and structures across five <italic>Sorbus</italic> species revealed extensive collinear regions, encompassing structural homology, translocations, and inversions (<xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7</bold>
</xref>). The longest sequence-similarity segments between species pairs were: 84,781 bp (<italic>M. alnifolia</italic> vs <italic>S. aucuparia</italic>), 97,089 bp (<italic>S. aucuparia</italic> vs <italic>S. aucuparia</italic> subsp. <italic>pohuashanensis</italic>), 229,635 bp (<italic>K. bristoliensis</italic> vs <italic>T. glaberrima</italic>), and 101,625 bp (<italic>T. glaberrima</italic> vs <italic>M. alnifolia</italic>). The highest degree of syntenic segment was observed between <italic>K. bristoliensis</italic> and <italic>T. glaberrima</italic> (310,482 bp), accounting for 80.28% of the <italic>K. bristoliensis</italic> mitochondrial genome length. Additionally, a 76,099 bp inversion structure was identified in the <italic>K. bristoliensis</italic> and <italic>T. glaberrima</italic> (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S10</bold>
</xref>).</p>
<fig id="f7" position="float">
<label>Figure&#xa0;7</label>
<caption>
<p>Collinearity Analysis of Mitochondrial Genomes in Five <italic>Sorbus</italic> s.l. Species. The gray arcs represent structurally homologous sequences, while other colors indicate structural rearrangements: inversions (green), transpositions (blue), and duplications (orange).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1619267-g007.tif">
<alt-text content-type="machine-generated">Sankey diagram showing genetic synteny, inversion, duplication, and translocation between five species: Karpatiosorbus bristoliensis, Torminalis glaberrima, Micromeles alnifolia, Sorbus aucuparia, and Sorbus aucuparia subsp. pohuashanensis. The flow lines indicate genetic relationships, with colors representing different genetic events: gray for synteny, green for inversion, blue for duplication, and orange for translocation.</alt-text>
</graphic>
</fig>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<title>Discussion</title>
<sec id="s4_1">
<title>Structure and size variation of chloroplast and mitochondrial genomes</title>
<p>This study reports the first complete mitochondrial and plastid genome assemblies for the endangered tree species <italic>K.&#xa0;bristoliensis</italic>. The circular mitochondrial genome spans 386,757 bp, demonstrating near-identical size conservation (1 bp difference) with its maternal progenitor <italic>Torminalis glaberrima</italic> (386,756 bp), consistent with its hybrid origin (<italic>Aria</italic> &#xd7; <italic>Torminalis</italic>). The chloroplast genome of <italic>K. bristoliensis</italic> is 68 bp smaller than that of <italic>Torminalis glaberrima</italic>. Compared with those of other species within <italic>Sorbus</italic> s.l., the mitochondrial genome of <italic>K. bristoliensis</italic> was smaller than those of <italic>M. alnifolia</italic> (45,5361 bp) and <italic>S. aucuparia</italic> subsp. <italic>pohuashanensis</italic> (396,857 bp) but larger than that of <italic>Sorbus aucuparia</italic> (384,977 bp). Previous studies have reported that the most minor mitochondrial genome in Rosaceae is <italic>Taihangia rupestris</italic> var. <italic>ciliata</italic> T.T.Yu &amp; C.L.Li, with a size of 265,633 bp (<xref ref-type="bibr" rid="B23">Li et&#xa0;al., 2025</xref>), and the largest one is <italic>Prunus mume</italic> (Siebold) Siebold &amp; Zucc. (535,727 bp) (<xref ref-type="bibr" rid="B57">Sun et&#xa0;al., 2022</xref>). The mitochondrial genome size of <italic>Sorbus</italic> s.s. falls within the range of mitochondrial genome size variation observed in Rosaceae. The mitochondrial genome of <italic>K. bristoliensis</italic> exhibited a GC content of 45.3%, aligning closely with the conserved genomic composition observed across Rosaceae mitogenomes (range: 43.31%&#x2013;45.62%); (<xref ref-type="bibr" rid="B32">Lu et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s4_2">
<title>Characteristics of repetitive sequences in the mitochondrial genome of <italic>Sorbus</italic> s.l.</title>
<p>SSRs exhibit high polymorphism and codominant inheritance, thus frequently used in population genetics studies (<xref ref-type="bibr" rid="B14">Hu et&#xa0;al., 2019</xref>). This study identified a large number of SSRs within the mitochondrial genome of <italic>Sorbus</italic> s.l., which exhibit potential utility for population genetic investigations. The number of SSRs in the mitochondrial genome of <italic>Sorbus</italic> s.l. is comparable to that observed in other Rosaceae species such as <italic>Rubus idaeus</italic> L (<xref ref-type="bibr" rid="B75">Zhang et&#xa0;al., 2025</xref>), yet demonstrates a lower frequency compared to members of the <italic>Malus</italic> genus within the same family (<xref ref-type="bibr" rid="B60">Wang et&#xa0;al., 2024</xref>). Consistent with previous findings, monomeric repeats predominated among the three SSR motif categories, exhibiting the highest numerical abundance (<xref ref-type="bibr" rid="B41">Niu et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B59">Sun et al., 2024</xref>). Tandem repeats may contribute to mitochondrial genome size expansion, particularly in species with exceptionally large genomes (<xref ref-type="bibr" rid="B67">Wynn and Christensen, 2019</xref>). Some tandem repeats exhibit species-specific amplification, serving as molecular markers for phylogenetic studies. A total of 107 tandem repeats were detected among the five species, which will be helpful for future studies. Dispersed repeats are nonadjacent homologous sequences ranging from tens to thousands of base pairs and are often distributed throughout the mitochondrial genome. These repeats facilitate homologous recombination, a key driver of structural rearrangements such as inversions, duplications, and the generation of subgenomic circles (<xref ref-type="bibr" rid="B65">Wu and Sloan, 2019</xref>). Previous studies have demonstrated prevalent mitochondrial genome rearrangements in Rosaceae species, as exemplified by members of the genus <italic>Malus</italic> (<xref ref-type="bibr" rid="B60">Wang et&#xa0;al., 2024</xref>) and <italic>Fragaria</italic> (<xref ref-type="bibr" rid="B9">Fan et&#xa0;al., 2022</xref>). Notably, one large palindromic repeat (26,130 bp) and forward repeat (24,730 bp) were detected in the mitochondrial genome of <italic>S. aucuparia</italic>. In addition, the <italic>S. aucuparia</italic> subsp. <italic>pohuashanensis</italic> mitogenome also presented one forward repeat exceeding 6,000 bp (6452 bp). Whether these large repeats are involved in mitochondrial genome rearrangement and other structural evolution remains to be further studied.</p>
</sec>
<sec id="s4_3">
<title>Gene transfer between mitochondrial and chloroplast genomes</title>
<p>Horizontal gene transfer (HGT) from chloroplasts to mitochondrial genomes represents a notable phenomenon in plant organelle evolution, reflecting complex inter-organellar genomic interactions (<xref ref-type="bibr" rid="B68">Yang et&#xa0;al., 2022</xref>). Although chloroplast-to-mitochondrion gene transfers occur less frequently than chloroplast-to-nuclear transfers do, accumulating evidence reveals their evolutionary significance in mitogenome evolution (<xref ref-type="bibr" rid="B13">Hong et&#xa0;al., 2021</xref>). In this study, we identified some chloroplast fragments transferred to the mitochondrial genome, with sizes ranging from 3,021 bp to 3,856 bp. Compared with previous studies in other Rosaceae, such as <italic>Rubus idaeus</italic> L. (46,456 bp transferred from chloroplasts to mitochondria) (<xref ref-type="bibr" rid="B75">Zhang et&#xa0;al., 2025</xref>), we found that homologous fragment transfer between the plastome and mitogenome in <italic>Sorbus</italic> s.l. was relatively rare. These findings indicate that the mitochondrial and chloroplast genomes of <italic>Sorbus</italic> s.l. are relatively highly conserved. Consistent with previous studies (<xref ref-type="bibr" rid="B61">Wang et&#xa0;al., 2007</xref>), we also found that the most frequently transferred genes were tRNAs, with a total of five tRNA genes (<italic>trnW-CCA, trnD-GUC, trnH-GUG, trnN-GUU, and trnM-CAU</italic>) shared by the plastid genome and the mitochondrial genome.</p>
</sec>
<sec id="s4_4">
<title>RNA editing in <italic>Sorbus</italic> s.l. mitochondrial genome</title>
<p>RNA editing is frequently reported in plant mitochondrial genomes (<xref ref-type="bibr" rid="B29">Liu et&#xa0;al., 2024</xref>). It is a crucial posttranscriptional modification process that plays a crucial role in controlling gene expression and functionality (<xref ref-type="bibr" rid="B27">Liu et&#xa0;al., 2020</xref>). At present, the common type of RNA editing is C&#x2013;U editing, and G-to-C and T-to-A editing can also occur (<xref ref-type="bibr" rid="B29">Liu et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B17">Ichinose and Sugita, 2016</xref>; <xref ref-type="bibr" rid="B68">Yang et&#xa0;al., 2022</xref>). In this study, all RNA editing sites in <italic>Sorbus</italic> s.l. were associated with C&#x2013;U editing. The number of RNA editing sites differed greatly in <italic>Sorbus</italic> s.l., ranging from 352 (<italic>T. glaberrima</italic>) to 463 (<italic>S. aucuparia subsp. pohuashanensis</italic>). Compared with those in other Rosaceae species, the number of RNA editing sites in <italic>Sorbus</italic> s.l. species was greater than that in <italic>Prunus pedunculata</italic> (262) (<xref ref-type="bibr" rid="B28">Liu et&#xa0;al., 2023</xref>) but less than that in <italic>Rubus chingii</italic> var. <italic>suavissimus</italic> (492) (<xref ref-type="bibr" rid="B54">Shi et&#xa0;al., 2024</xref>) and <italic>Taihangia rupestris</italic> var. <italic>ciliata</italic> (470) (<xref ref-type="bibr" rid="B23">Li et&#xa0;al., 2025</xref>). Overall, RNA editing sites exhibit significant variations&#xa0;across different species and even within the same genus, such as <italic>S. aucuparia</italic> subsp. <italic>pohuashanensis</italic> and <italic>S.&#xa0;aucuparia</italic>. Notably, the RNA editing sites appear to exhibit a gene preference, with editing events occurring most frequently within the NADH dehydrogenase genes. Similar results have been reported in other species, such as <italic>Prunus pedunculata</italic> (Pall.) Maxim. (<xref ref-type="bibr" rid="B28">Liu et&#xa0;al., 2023</xref>) and <italic>Gelsemium elegans</italic> (Gardner &amp; Champ.) Benth (<xref ref-type="bibr" rid="B69">You et&#xa0;al., 2023</xref>). We also observed that RNA editing influenced the start and termination codons of protein-coding genes. For example, the start codons of <italic>cox1, nad1, rpl16, and rps4</italic> changed from ACG to AUG through RNA editing, whereas RNA editing led to the termination codons of <italic>atp6</italic>. RNA editing events that alter initiation and termination codons are frequently observed, as exemplified by modifications to the <italic>nad1</italic> initiation codon and <italic>ccmFc</italic> termination codon in the mitochondrial genome of <italic>Garcinia mangostana</italic> L (<xref ref-type="bibr" rid="B62">Wee et&#xa0;al., 2022</xref>).</p>
</sec>
<sec id="s4_5">
<title>Capture of polymorphic loci</title>
<p>The mitochondrial and chloroplast genomes harbor a substantial number of polymorphic sequences that can be developed as DNA barcodes for phylogenetic and population genetic studies (<xref ref-type="bibr" rid="B24">Li and Wei, 2022</xref>). This study analyzed the mitochondrial genomes of five <italic>Sorbus</italic> s.l. species and revealed that the <italic>nad4</italic> gene presented the greatest number of nucleotide polymorphisms. These findings suggest its potential utility as an effective DNA barcode for investigating phylogenetic relationships within <italic>Sorbus</italic> s.l. Through comparative analysis of chloroplast genomes, we identified six plastid gene regions exhibiting high genetic polymorphism, which could serve as chloroplast DNA barcodes for investigating phylogenetic relationships within <italic>Sorbus</italic> s.l. Furthermore, the <italic>ycf1</italic> gene, conventionally employed as a universal barcode for angiosperms (<xref ref-type="bibr" rid="B7">Dong et&#xa0;al., 2015</xref>), demonstrated lower nucleotide polymorphism in this study.</p>
</sec>
<sec id="s4_6">
<title>Maternal lineage phylogenetic analysis of the <italic>Sorbus</italic> s.l.</title>
<p>In this study, we provide a high-resolution maternal genetic framework for <italic>Sorbus</italic> s.l. <italic>Aria pannonica</italic> (K&#xe1;rp&#xe1;ti) Sennikov &amp; Kurtto, which is a hybrid-origin species with a ploidy level of triploid (<xref ref-type="bibr" rid="B56">Somlyay and Sennikov, 2015</xref>). <italic>A. pannonica</italic> originated from intragenus hybridization because of its morphological similarity to <italic>A. nivea</italic> Host and <italic>A. graeca</italic> (Spach) M.Roem (<xref ref-type="bibr" rid="B56">Somlyay and Sennikov, 2015</xref>). However, <italic>A. pannonica</italic> is more closely related to the genus <italic>Chamaemespilus</italic> in our plastome-based phylogeny. Therefore, we inferred that the female parent of this hybrid species may have originated from <italic>Chamaemespilus</italic>. This fact indicated that <italic>Aria pannonica</italic> should treat as the member of <italic>Majovskya</italic>. The genus <italic>Karpatiosorbus</italic> is thought to originate from the hybridization of <italic>Aria</italic> and <italic>Torminalis</italic> (<xref ref-type="bibr" rid="B42">Pellicer et&#xa0;al., 2012</xref>). Both our mitogenome- and platome-based phylogenetic data supported that <italic>Karpatiosorbus</italic> and <italic>Torminalis</italic> clustered into one branch. These findings indicate that the female parent of this genus is <italic>Torminalis</italic>. The genus <italic>Hedlundia</italic> contains one sexual hybrid and 39 apomictic species and originates from crosses between various species of <italic>Aria</italic> and <italic>Sorbus</italic> s.l (<xref ref-type="bibr" rid="B52">Sennikov and Kurtto, 2017</xref>). The genus <italic>Hedlundia</italic> is separated into two clades: one clade is closely related to <italic>Micromeles</italic>, and the other clade is embedded in <italic>Sorbus</italic> s.s, indicating the complex origin of the hybrid genus. <xref ref-type="bibr" rid="B52">Sennikov and Kurtto (2017)</xref> proposed the establishment of the new nothogenus <italic>Sorbomeles</italic> to accommodate hybrids of <italic>Sorbus</italic> and <italic>Micromeles</italic> origin. In the present study, molecular evidence suggests <italic>Micromeles</italic> likely served as the maternal progenitor of <italic>Hedlundia austriaca</italic> (Beck) Sennikov &amp; Kurtto and <italic>H. persica</italic> (Hedl.) Mezhenskyj. Consequently, these two taxa are hereby formally proposed for reclassification into the hybrid genus &#xd7;<italic>Sorbomeles.</italic> A previous study indicated that hybridization between <italic>Aria</italic> and <italic>Sorbus</italic> s.s. contributed to the genus <italic>Micromeles</italic>. Both our results and those of previous plastome-based studies support that <italic>Micromeles</italic> is a sister to <italic>Sorbus</italic> s.s., whereas this is inconsistent with nuclear phylogenetic analyses (<xref ref-type="bibr" rid="B73">Zhang et&#xa0;al., 2023</xref>). Therefore, hybridization occurs between <italic>Micromeles</italic> and <italic>Sorbus</italic> s.s. <italic>Scandosorbus intermedia</italic> (Ehrh.) Sennikov originates from hybridization among <italic>Sorbus</italic> s.s., <italic>Aria</italic>, and <italic>Torminalis</italic> (<xref ref-type="bibr" rid="B42">Pellicer et&#xa0;al., 2012</xref>). It involves a complex species formation process. In this study, <italic>S. intermedia</italic> was a sister to <italic>S. dumosa</italic>. These findings indicate that the female parent of this species is <italic>S. dumosa</italic> House or a related species.</p>
<p>Phylogenetic analyses have consistently recovered <italic>Sorbus</italic> s.s. as a monophyletic clade distinct from other simple-leaved genera and the compound-leaved genus <italic>Cormus</italic> in previous studies (<xref ref-type="bibr" rid="B5">Campbell et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B76">Zheng and Zhang, 2007</xref>; <xref ref-type="bibr" rid="B44">Potter et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B30">Lo and Donoghue, 2012</xref>). This limitation arises from inadequate sampling in prior studies, which excluded phylogenetically critical hybrid-origin taxa. However, our current phylogenomic reconstruction reveals that <italic>Hedlundia</italic> and <italic>Scandosorbus</italic> (simple-leaved hybrid-origin genera) nests within <italic>Sorbus</italic> s.s., thereby rendering the latter non-monophyletic. This phylogenetic incongruence results from the hybrid origin of <italic>Hedlundia</italic> and <italic>Scandosorbus</italic>, which inherited its maternal genome from ancestral <italic>Sorbus</italic> s.s. lineages. <italic>Sorbus</italic> s.s. is one of the most typical examples of taxonomic complexity arising from the combined effects of hybridization, polyploidy and apomixis in the Rosaceae (<xref ref-type="bibr" rid="B49">Robertson et&#xa0;al., 2010</xref>). Previous research has demonstrated that both insect-pollinated and bird-pollinated are present in the genus <italic>Sorbus</italic> s.s.; moreover, mammals also play an important role in the spread of <italic>Sorbus</italic> s.s. seeds (<xref ref-type="bibr" rid="B45">Rasp&#xe9; et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B1">Bednorz et&#xa0;al., 2006</xref>). Some species of this genus are self-incompatible (<xref ref-type="bibr" rid="B45">Rasp&#xe9; et&#xa0;al., 2000</xref>). These factors strongly increase the frequency of interspecific hybridization in <italic>Sorbus</italic> s.s. as well as hybrid with other genera. Therefore, elucidating taxonomic challenges in <italic>Sorbus</italic> s.s requires resolving hybridization patterns and clarifying phylogenetic placements of hybrid-origin taxa within the genus.</p>
</sec>
<sec id="s4_7">
<title>Frequent structural rearrangements of mitogenome</title>
<p>Mitochondrial genome structural complexity arises from extensive repetitive sequences and frequent structural rearrangements. Such mitochondrial genome rearrangements have also been frequently reported in other genera of Rosaceae, notably <italic>Prunus</italic> (<xref ref-type="bibr" rid="B71">Zhai et&#xa0;al., 2025</xref>) and <italic>Malus</italic> (<xref ref-type="bibr" rid="B60">Wang et&#xa0;al., 2024</xref>). This study revealed that <italic>K. bristoliensis</italic> shares substantial homologous segments with <italic>T. glaberrima</italic> (species from its maternal genus), while exhibiting limited homology with phylogenetically distant genera. This pattern aligns with their close phylogenetic origin and the hybrid origin of <italic>Karpatiosorbus</italic>. Despite the extensive shared homology, a 76,099 bp inversion structure was identified between their mitochondrial genomes, suggesting incipient mitogenomic divergence between the two genera.</p>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<title>Conclusions</title>
<p>In this study, we reported the first complete mitochondrial and plastid genomes of the endangered tree <italic>Karpatiosorbus bristoliensis</italic> utilizing PacBio-HiFi long reads. <italic>K. bristoliensis</italic> mitochondrial and plastid genomes consists of one circular chromosome structures with the length of 386,757 bp bp and 160,322 bp, respectively. We analyzed repeat sequences, RNA editing, inter-organellar gene transfer, and pi value in the mitochondrial genome of <italic>Sorbus</italic> s.l., enriching our knowledge of mitochondrial genome evolution of this genus. We also conducted phylogenetic analyses within <italic>Sorbus</italic> s.l. utilizing mitochondrial and plastid genome. We emphasize that resolving hybridization dynamics in <italic>Sorbus</italic> s.l. is critical for achieving a taxonomically coherent delineation of the genus. Overall, our study provided a high-resolution maternal framework for <italic>Sorbus</italic> s.l.</p>
</sec>
</body>
<back>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The mitochondrial and plastid genome have been deposited in GenBank under the accession numbers: PV207325 and PV207326.</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>QL: Funding acquisition, Conceptualization, Investigation, Supervision, Writing &#x2013; review &amp; editing, Resources, Software, Writing &#x2013; original draft, Project administration, Validation, Formal Analysis, Data curation, Methodology, Visualization. RW: Methodology, Writing &#x2013; review &amp; editing, Software, Visualization.</p>
</sec>
<sec id="s8" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare that no financial support was received for the research and/or publication of this article.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>We are grateful to Dr Maarten J.M. Christenhusz for providing the raw data of <italic>Karpatiosorbus bristoliensis</italic>.</p>
</ack>
<sec id="s9" 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="s10" 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="s11" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s12" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fpls.2025.1619267/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fpls.2025.1619267/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Image1.tif" id="SF1" mimetype="image/tiff">
<label>Supplementary Figure&#xa0;1</label>
<caption>
<p>Bandage visual mitochondrial genome assembly map.</p>
</caption>
</supplementary-material>
<supplementary-material xlink:href="Table1.xlsx" id="SM1" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="Table2.xlsx" id="SM2" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="Table3.xlsx" id="SM3" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="Table4.xlsx" id="SM4" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="Table5.xlsx" id="SM5" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="Table6.xlsx" id="SM6" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="Table7.xlsx" id="SM7" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="Table8.xlsx" id="SM8" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="Table9.xlsx" id="SM9" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="Table10.xlsx" id="SM10" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
</sec>
<ref-list>
<title>References</title>
<ref id="B1">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Bednorz</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Myczko</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Kosi&#x144;ski</surname> <given-names>P.</given-names>
</name>
</person-group> (<year>2006</year>). <article-title>Genetic variability and structure of the wild service tree (<italic>Sorbus torminalis</italic> (L.) crantz) in Poland</article-title>. <source>Silvae Genet.</source> <volume>55</volume>, <fpage>197</fpage>&#x2013;<lpage>202</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1515/sg-2006-0027</pub-id>
</citation></ref>
<ref id="B2">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Benson</surname> <given-names>G.</given-names>
</name>
</person-group> (<year>1999</year>). <article-title>Tandem repeats finder: a program to analyze DNA sequences</article-title>. <source>Nucleic Acids Res.</source> <volume>27</volume>, <fpage>573</fpage>&#x2013;<lpage>580</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/nar/27.2.573</pub-id>, PMID: <pub-id pub-id-type="pmid">9862982</pub-id></citation></ref>
<ref id="B3">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Bi</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Qu</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Hou</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Wu</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Ye</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Yin</surname> <given-names>T.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>Deciphering the multi-chromosomal mitochondrial genome of <italic>populus simonii</italic>
</article-title>. <source>Front. Plant Sci.</source> <volume>13</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fpls.2022.914635</pub-id>, PMID: <pub-id pub-id-type="pmid">35783945</pub-id></citation></ref>
<ref id="B4">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Bi</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Shen</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Han</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Qu</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Hou</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Xu</surname> <given-names>K.</given-names>
</name>
<etal/>
</person-group>. (<year>2024</year>). <article-title>PMAT: an efficient plant mitogenome assembly toolkit using low-coverage HiFi sequencing data</article-title>. <source>Hortic. Res.</source> <volume>11</volume>, <elocation-id>uhae023</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/hr/uhae023</pub-id>, PMID: <pub-id pub-id-type="pmid">38469379</pub-id></citation></ref>
<ref id="B5">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Campbell</surname> <given-names>C. S.</given-names>
</name>
<name>
<surname>Evans</surname> <given-names>R. C.</given-names>
</name>
<name>
<surname>Morgan</surname> <given-names>D. R.</given-names>
</name>
<name>
<surname>Dickinson</surname> <given-names>T. A.</given-names>
</name>
<name>
<surname>Arsenault</surname> <given-names>M. P.</given-names>
</name>
</person-group> (<year>2007</year>). <article-title>Phylogeny of subtribe Pyrinae (formerly the Maloideae, Rosaceae): Limited resolution of a complex evolutionary history</article-title>. <source>Plant Syst. Evol.</source> <volume>266</volume>, <fpage>119</fpage>&#x2013;<lpage>145</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s00606-007-0545-y</pub-id>
</citation></ref>
<ref id="B6">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Chester</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Cowan</surname> <given-names>R. S.</given-names>
</name>
<name>
<surname>Fay</surname> <given-names>M. F.</given-names>
</name>
<name>
<surname>Rich</surname> <given-names>T. C. G.</given-names>
</name>
</person-group> (<year>2007</year>). <article-title>Parentage of endemic <italic>Sorbus</italic> L. (Rosaceae) species in the British Isles: Evidence from plastid DNA</article-title>. <source>Bot. J. Linn. Soc</source> <volume>154</volume>, <fpage>291</fpage>&#x2013;<lpage>304</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/j.1095-8339.2007.00669.x</pub-id>
</citation></ref>
<ref id="B7">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Dong</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Xu</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Sun</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Zuo</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Shi</surname> <given-names>S.</given-names>
</name>
<etal/>
</person-group>. (<year>2015</year>). <article-title>
<italic>ycf1</italic>, the most promising plastid DNA barcode of land plants</article-title>. <source>Sci. Rep.</source> <volume>5</volume>, <elocation-id>8348</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/srep08348</pub-id>, PMID: <pub-id pub-id-type="pmid">25672218</pub-id></citation></ref>
<ref id="B8">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Edera</surname> <given-names>A. A.</given-names>
</name>
<name>
<surname>Small</surname> <given-names>I.</given-names>
</name>
<name>
<surname>Milone</surname> <given-names>D. H.</given-names>
</name>
<name>
<surname>Sanchez-Puerta</surname> <given-names>M. V.</given-names>
</name>
</person-group> (<year>2021</year>). <article-title>Deepred-Mt: Deep representation learning for predicting C-to-U RNA editing in plant mitochondria</article-title>. <source>Comput. Biol. Med.</source> <volume>136</volume>, <elocation-id>104682</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.compbiomed.2021.104682</pub-id>, PMID: <pub-id pub-id-type="pmid">34343887</pub-id></citation></ref>
<ref id="B9">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Fan</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Liu</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Jia</surname> <given-names>Q.</given-names>
</name>
<name>
<surname>Du</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Chen</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Ruan</surname> <given-names>J.</given-names>
</name>
<etal/>
</person-group>. (<year>2022</year>). <article-title>
<italic>Fragaria</italic> mitogenomes evolve rapidly in structure but slowly in sequence and incur frequent multinucleotide mutations mediated by microinversions</article-title>. <source>New Phytol.</source> <volume>236</volume>, <fpage>745</fpage>&#x2013;<lpage>759</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/nph.18334</pub-id>, PMID: <pub-id pub-id-type="pmid">35731093</pub-id></citation></ref>
<ref id="B10">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Goel</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Schneeberger</surname> <given-names>K.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>plotsr: visualizing structural similarities and rearrangements between multiple genomes</article-title>. <source>Bioinformatics</source> <volume>38</volume>, <fpage>2922</fpage>&#x2013;<lpage>2926</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/bioinformatics/btac196</pub-id>, PMID: <pub-id pub-id-type="pmid">35561173</pub-id></citation></ref>
<ref id="B11">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Goel</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Sun</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Jiao</surname> <given-names>W.-B.</given-names>
</name>
<name>
<surname>Schneeberger</surname> <given-names>K.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>SyRI: finding genomic rearrangements and local sequence differences from whole-genome assemblies</article-title>. <source>Genome Biol.</source> <volume>20</volume>, <fpage>277</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1186/s13059-019-1911-0</pub-id>, PMID: <pub-id pub-id-type="pmid">31842948</pub-id></citation></ref>
<ref id="B12">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Gualberto</surname> <given-names>J. M.</given-names>
</name>
<name>
<surname>Newton</surname> <given-names>K. J.</given-names>
</name>
</person-group> (<year>2017</year>). <article-title>Plant mitochondrial genomes: dynamics and mechanisms of mutation</article-title>. <source>Annu. Rev. Plant Biol.</source> <volume>68</volume>, <fpage>225</fpage>&#x2013;<lpage>252</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1146/annurev-arplant-043015-112232</pub-id>, PMID: <pub-id pub-id-type="pmid">28226235</pub-id></citation></ref>
<ref id="B13">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Hong</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Liao</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Ye</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Yang</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Zhu</surname> <given-names>W.</given-names>
</name>
<etal/>
</person-group>. (<year>2021</year>). <article-title>A complete mitochondrial genome for fragrant Chinese rosewood (<italic>Dalbergia odorifera</italic>, Fabaceae) with comparative analyses of genome structure and intergenomic sequence transfers</article-title>. <source>BMC Genomics</source> <volume>22</volume>, <fpage>672</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1186/s12864-021-07967-7</pub-id>, PMID: <pub-id pub-id-type="pmid">34536995</pub-id></citation></ref>
<ref id="B14">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Hu</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Zhou</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Chen</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Hu</surname> <given-names>X.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>Advancing phylogeography with chloroplast DNA markers</article-title>. <source>Biodivers. Sci.</source> <volume>27</volume>, <fpage>219</fpage>&#x2013;<lpage>234</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.17520/biods.2018319</pub-id>
</citation></ref>
<ref id="B15">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Huang</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Xu</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Hu</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Jiang</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Sun</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Zhao</surname> <given-names>W.</given-names>
</name>
<etal/>
</person-group>. (<year>2024</year>a). <article-title>The Mitochondrial Genome of <italic>Cathaya argyrophylla</italic> Reaches 18. 99 Mb: Analysis of Super-Large Mitochondrial Genomes in Pinaceae</article-title>. <source>arXiv</source>. Available online at: <uri xlink:href="https://arxiv.org/pdf/2410.07006">https://arxiv.org/pdf/2410.07006</uri>.</citation></ref>
<ref id="B16">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Huang</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Yu</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Xu</surname> <given-names>W.</given-names>
</name>
</person-group> (<year>2024</year>b). <article-title>CPStools: A package for analyzing chloroplast genome sequences</article-title>. <source>iMetaOmics</source> <volume>1</volume>, <elocation-id>e25</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1002/imo2.25</pub-id>
</citation></ref>
<ref id="B17">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ichinose</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Sugita</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2016</year>). <article-title>RNA editing and its molecular mechanism in plant organelles</article-title>. <source>Genes (Basel).</source> <volume>8</volume>, <elocation-id>5</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/genes8010005</pub-id>, PMID: <pub-id pub-id-type="pmid">28025543</pub-id></citation></ref>
<ref id="B18">
<citation citation-type="book">
<person-group person-group-type="author">
<collab>IUCN</collab>
</person-group> (<year>2001</year>). <source>IUCN Red List Categories and Criteria: Version 3.1</source> (<publisher-loc>Gland, Switzerland</publisher-loc>: <publisher-name>IUCN Species Survival Commission</publisher-name>). Available online at: <uri xlink:href="https://portals.iucn.org/library/sites/library/files/documents/RL-2001-001.pdf">https://portals.iucn.org/library/sites/library/files/documents/RL-2001-001.pdf</uri> (Accessed <access-date>February 9, 2000</access-date>).</citation></ref>
<ref id="B19">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Kalyaanamoorthy</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Minh</surname> <given-names>B. Q.</given-names>
</name>
<name>
<surname>Wong</surname> <given-names>T. K. F.</given-names>
</name>
<name>
<surname>von Haeseler</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Jermiin</surname> <given-names>L. S.</given-names>
</name>
</person-group> (<year>2017</year>). <article-title>ModelFinder: fast model selection for accurate phylogenetic estimates</article-title>. <source>Nat. Methods</source> <volume>14</volume>, <fpage>587</fpage>&#x2013;<lpage>589</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/nmeth.4285</pub-id>, PMID: <pub-id pub-id-type="pmid">28481363</pub-id></citation></ref>
<ref id="B20">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Katoh</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Rozewicki</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Yamada</surname> <given-names>K. D.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>MAFFT online service: Multiple sequence alignment, interactive sequence choice and visualization</article-title>. <source>Brief. Bioinform.</source> <volume>20</volume>, <fpage>1160</fpage>&#x2013;<lpage>1166</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/bib/bbx108</pub-id>, PMID: <pub-id pub-id-type="pmid">28968734</pub-id></citation></ref>
<ref id="B21">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Kurtz</surname> <given-names>S.</given-names>
</name>
</person-group> (<year>2001</year>). <article-title>REPuter: the manifold applications of repeat analysis on a genomic scale</article-title>. <source>Nucleic Acids Res.</source> <volume>29</volume>, <fpage>4633</fpage>&#x2013;<lpage>4642</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/nar/29.22.4633</pub-id>, PMID: <pub-id pub-id-type="pmid">11713313</pub-id></citation></ref>
<ref id="B22">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Li</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Ni</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Lu</surname> <given-names>Q.</given-names>
</name>
<name>
<surname>Chen</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Liu</surname> <given-names>C.</given-names>
</name>
</person-group> (<year>2024</year>). <article-title>PMGA: A plant mitochondrial genome annotator</article-title>. <source>Plant Commun.</source> <volume>6</volume>, <elocation-id>101191</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.xplc.2024.101191</pub-id>, PMID: <pub-id pub-id-type="pmid">39521957</pub-id></citation></ref>
<ref id="B23">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Li</surname> <given-names>Z.-Z.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>He</surname> <given-names>X.-Y.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>W.-G.</given-names>
</name>
</person-group> (<year>2025</year>). <article-title>The <italic>Taihangia</italic> mitogenome provides new insights into its adaptation and organelle genome evolution in Rosaceae</article-title>. <source>Planta</source> <volume>261</volume>, <fpage>59</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s00425-025-04629-w</pub-id>, PMID: <pub-id pub-id-type="pmid">39939538</pub-id></citation></ref>
<ref id="B24">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Li</surname> <given-names>Q.</given-names>
</name>
<name>
<surname>Wei</surname> <given-names>R.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>Comparison of boraginales plastomes: insights into codon usage bias, adaptive evolution, and phylogenetic relationships</article-title>. <source>Diversity</source> <volume>14</volume>, <elocation-id>1104</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/d14121104</pub-id>
</citation></ref>
<ref id="B25">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lin</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Shao</surname> <given-names>B.</given-names>
</name>
<name>
<surname>Gao</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>T.</given-names>
</name>
<etal/>
</person-group>. (<year>2025</year>). <article-title>Phylogenomics of angiosperms based on mitochondrial genes: insights into deep node relationships</article-title>. <source>BMC Biol.</source> <volume>23</volume>, <fpage>45</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1186/s12915-025-02135-9</pub-id>, PMID: <pub-id pub-id-type="pmid">39948594</pub-id></citation></ref>
<ref id="B26">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Linnaeus</surname> <given-names>C.</given-names>
</name>
</person-group> (<year>1753</year>). <source>Species plantarum</source> Vol. <volume>2</volume> (<publisher-loc>Stockholm</publisher-loc>: <publisher-name>impensis Laurentii Salvii</publisher-name>). Holmiae. doi:&#xa0;<pub-id pub-id-type="doi">10.5962/bhl.title.669</pub-id>
</citation></ref>
<ref id="B27">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Liu</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Cao</surname> <given-names>S.-K.</given-names>
</name>
<name>
<surname>Sayyed</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Yang</surname> <given-names>H.-H.</given-names>
</name>
<name>
<surname>Zhao</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>X.</given-names>
</name>
<etal/>
</person-group>. (<year>2020</year>). <article-title>The DYW-subgroup pentatricopeptide repeat protein PPR27 interacts with ZmMORF1 to facilitate mitochondrial RNA editing and seed development in maize</article-title>. <source>J. Exp. Bot.</source> <volume>71</volume>, <fpage>5495</fpage>&#x2013;<lpage>5505</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/jxb/eraa273</pub-id>, PMID: <pub-id pub-id-type="pmid">32531050</pub-id></citation></ref>
<ref id="B28">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Liu</surname> <given-names>Q.</given-names>
</name>
<name>
<surname>Wu</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Tian</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Yang</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Liu</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Feng</surname> <given-names>Y.</given-names>
</name>
<etal/>
</person-group>. (<year>2023</year>). <article-title>Complete mitochondrial genome of the endangered <italic>Prunus pedunculata</italic> (Prunoideae, Rosaceae) in China: characterization and phylogenetic analysis</article-title>. <source>Front. Plant Sci.</source> <volume>14</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fpls.2023.1266797</pub-id>, PMID: <pub-id pub-id-type="pmid">38155854</pub-id></citation></ref>
<ref id="B29">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Liu</surname> <given-names>G. H.</given-names>
</name>
<name>
<surname>Zuo</surname> <given-names>Y. W.</given-names>
</name>
<name>
<surname>Shan</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Yu</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>J. X.</given-names>
</name>
<name>
<surname>Chen</surname> <given-names>Y.</given-names>
</name>
<etal/>
</person-group>. (<year>2024</year>). <article-title>Structural analysis of the mitochondrial genome of Santalum album reveals a complex branched configuration</article-title>. <source>Genomics</source> <volume>116</volume>, <elocation-id>110935</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.ygeno.2024.110935</pub-id>, PMID: <pub-id pub-id-type="pmid">39243912</pub-id></citation></ref>
<ref id="B30">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lo</surname> <given-names>E. Y. Y.</given-names>
</name>
<name>
<surname>Donoghue</surname> <given-names>M. J.</given-names>
</name>
</person-group> (<year>2012</year>). <article-title>Expanded phylogenetic and dating analyses of the apples and their relatives (Pyreae, Rosaceae)</article-title>. <source>Mol. Phylogenet. Evol.</source> <volume>63</volume>, <fpage>230</fpage>&#x2013;<lpage>243</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.ympev.2011.10.005</pub-id>, PMID: <pub-id pub-id-type="pmid">22293154</pub-id></citation></ref>
<ref id="B31">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Lu</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Stephen</surname> <given-names>A. ,. S.</given-names>
</name>
</person-group> (<year>2003</year>). <article-title>
<italic>Sorbus</italic> L</article-title>. Eds. <person-group person-group-type="editor">
<name>
<surname>Wu</surname> <given-names>Z. Y.</given-names>
</name>
<name>
<surname>Raven</surname> <given-names>P. H.</given-names>
</name>
<name>
<surname>Hong</surname> <given-names>D. Y.</given-names>
</name>
</person-group>
<source>Flora of China</source> (<publisher-loc>SaintLouis</publisher-loc>: <publisher-name>Science Press, Beijing &amp; Missouri Botanical Garden Press</publisher-name>), <fpage>144</fpage>&#x2013;<lpage>170</lpage>.</citation></ref>
<ref id="B32">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lu</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Que</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>Y.</given-names>
</name>
</person-group> (<year>2023</year>). <article-title>Assembly and analysis of the first complete mitochondrial genome of <italic>Punica granatum</italic> and the gene transfer from chloroplast genome</article-title>. <source>Front. Plant Sci.</source> <volume>14</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fpls.2023.1132551</pub-id>, PMID: <pub-id pub-id-type="pmid">37416882</pub-id></citation></ref>
<ref id="B33">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lu</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Wu</surname> <given-names>Q.</given-names>
</name>
<name>
<surname>Sun</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Yang</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Wei</surname> <given-names>E.</given-names>
</name>
<etal/>
</person-group>. (<year>2025</year>). <article-title>Multiple chromosomal configurations and phylogenetic implications in Saccharum</article-title>. <source>J. Integr. Agric</source>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jia.2025.02.018</pub-id>
</citation></ref>
<ref id="B34">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Mar&#xe7;ais</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Delcher</surname> <given-names>A. L.</given-names>
</name>
<name>
<surname>Phillippy</surname> <given-names>A. M.</given-names>
</name>
<name>
<surname>Coston</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Salzberg</surname> <given-names>S. L.</given-names>
</name>
<name>
<surname>Zimin</surname> <given-names>A.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>MUMmer4: A fast and versatile genome alignment system</article-title>. <source>PloS Comput. Biol.</source> <volume>14</volume>, <elocation-id>e1005944</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1371/journal.pcbi.1005944</pub-id>, PMID: <pub-id pub-id-type="pmid">29373581</pub-id></citation></ref>
<ref id="B35">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>McAllister</surname> <given-names>H.</given-names>
</name>
</person-group> (<year>2005</year>). <source>The genus Sorbus Mountain Ash and other Rowans</source> (<publisher-loc>London</publisher-loc>: <publisher-name>Royal Botanical Gardens</publisher-name>).</citation></ref>
<ref id="B36">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Mezhenska</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Mezhenskyj</surname> <given-names>V.</given-names>
</name>
<name>
<surname>Yakubenko</surname> <given-names>B.</given-names>
</name>
</person-group> (<year>2018</year>). <source>NULESU Collections of Fruit and Ornamental Plants</source> (<publisher-loc>Kiev</publisher-loc>: <publisher-name>Lira-K</publisher-name>). &#x41a;&#x41e;&#x41b;&#x415;&#x41a;&#x426;&#x406;&#x42f; &#x41d;&#x423;&#x411;&#x406;&#x41f; &#x423;&#x41a;&#x420;&#x410;&#x407;&#x41d;&#x418; &#x41f;&#x41b;&#x41e;&#x414;&#x41e;&#x412;&#x418;&#x425; &#x406; &#x414;&#x415;&#x41a;&#x41e;&#x420;&#x410;&#x422;&#x418;&#x412;&#x41d;&#x418;&#x425; &#x420;&#x41e;&#x421;&#x41b;&#x418;&#x41d;.</citation></ref>
<ref id="B37">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Minh</surname> <given-names>B. Q.</given-names>
</name>
<name>
<surname>Schmidt</surname> <given-names>H. A.</given-names>
</name>
<name>
<surname>Chernomor</surname> <given-names>O.</given-names>
</name>
<name>
<surname>Schrempf</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Woodhams</surname> <given-names>M. D.</given-names>
</name>
<name>
<surname>Von Haeseler</surname> <given-names>A.</given-names>
</name>
<etal/>
</person-group>. (<year>2020</year>). <article-title>IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era</article-title>. <source>Mol. Biol. Evol.</source> <volume>37</volume>, <fpage>1530</fpage>&#x2013;<lpage>1534</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/molbev/msaa015</pub-id>, PMID: <pub-id pub-id-type="pmid">32011700</pub-id></citation></ref>
<ref id="B38">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Morgante</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Hanafey</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Powell</surname> <given-names>W.</given-names>
</name>
</person-group> (<year>2002</year>). <article-title>Microsatellites are preferentially associated with nonrepetitive DNA in plant genomes</article-title>. <source>Nat. Genet.</source> <volume>30</volume>, <fpage>194</fpage>&#x2013;<lpage>200</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/ng822</pub-id>, PMID: <pub-id pub-id-type="pmid">11799393</pub-id></citation></ref>
<ref id="B39">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Nelson-Jones</surname> <given-names>E. B.</given-names>
</name>
<name>
<surname>Briggs</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Smith</surname> <given-names>A. G.</given-names>
</name>
</person-group> (<year>2002</year>). <article-title>The origin of intermediate species of the genus <italic>Sorbus</italic>
</article-title>. <source>Theor. Appl. Genet.</source> <volume>105</volume>, <fpage>953</fpage>&#x2013;<lpage>963</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s00122-002-0957-6</pub-id>, PMID: <pub-id pub-id-type="pmid">12582921</pub-id></citation></ref>
<ref id="B40">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>N&#xe9;meth</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Papp</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Noskov&#xe1;</surname> <given-names>J.</given-names>
</name>
<name>
<surname>H&#xf6;hn</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2020</year>). <article-title>Speciation by triparental hybridization in genus <italic>Sorbus</italic> (Rosaceae)</article-title>. <source>Biol. Futur.</source> <volume>71</volume>, <fpage>209</fpage>&#x2013;<lpage>222</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s42977-020-00003-x</pub-id>, PMID: <pub-id pub-id-type="pmid">34554510</pub-id></citation></ref>
<ref id="B41">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Niu</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Gao</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Liu</surname> <given-names>J.</given-names>
</name>
</person-group> (<year>2024</year>). <article-title>Mitochondrial genome variation and intergenomic sequence transfers in <italic>Hevea</italic> species</article-title>. <source>Front. Plant Sci.</source> <volume>15</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fpls.2024.1234643</pub-id>, PMID: <pub-id pub-id-type="pmid">38660449</pub-id></citation></ref>
<ref id="B42">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Pellicer</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Clermont</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Houston</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Rich</surname> <given-names>T. C. G.</given-names>
</name>
<name>
<surname>Fay</surname> <given-names>M. F.</given-names>
</name>
</person-group> (<year>2012</year>). <article-title>Cytotype diversity in the <italic>Sorbus</italic> complex (Rosaceae) in Britain: Sorting out the puzzle</article-title>. <source>Ann. Bot.</source> <volume>110</volume>, <fpage>1185</fpage>&#x2013;<lpage>1193</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/aob/mcs185</pub-id>, PMID: <pub-id pub-id-type="pmid">22922587</pub-id></citation></ref>
<ref id="B43">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Phipps</surname> <given-names>J. B.</given-names>
</name>
<name>
<surname>Robertson</surname> <given-names>K. R.</given-names>
</name>
<name>
<surname>Smith</surname> <given-names>P. G.</given-names>
</name>
<name>
<surname>Rohrer</surname> <given-names>J. R.</given-names>
</name>
</person-group> (<year>1990</year>). <article-title>A checklist of the subfamily Maloideae (Rosaceae)</article-title>. <source>Can. J. Bot.</source> <volume>68</volume>, <fpage>2209</fpage>&#x2013;<lpage>2269</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1139/b90-288</pub-id>
</citation></ref>
<ref id="B44">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Potter</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Eriksson</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Evans</surname> <given-names>R. C.</given-names>
</name>
<name>
<surname>Oh</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Smedmark</surname> <given-names>J. E. E.</given-names>
</name>
<name>
<surname>Morgan</surname> <given-names>D. R.</given-names>
</name>
<etal/>
</person-group>. (<year>2007</year>). <article-title>Phylogeny and classification of rosaceae</article-title>. <source>Plant Syst. Evol.</source> <volume>266</volume>, <fpage>5</fpage>&#x2013;<lpage>43</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s00606-007-0539-9</pub-id>
</citation></ref>
<ref id="B45">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rasp&#xe9;</surname> <given-names>O.</given-names>
</name>
<name>
<surname>Saumitou-Laprade</surname> <given-names>P.</given-names>
</name>
<name>
<surname>Cuguen</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Jacquemart</surname> <given-names>A. L.</given-names>
</name>
</person-group> (<year>2000</year>). <article-title>Chloroplast DNA haplotype variation and population differentiation in <italic>Sorbus aucuparia</italic> L. (Rosaceae: Maloideae)</article-title>. <source>Mol. Ecol.</source> <volume>9</volume>, <fpage>1113</fpage>&#x2013;<lpage>1122</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1046/j.1365-294X.2000.00977.x</pub-id>, PMID: <pub-id pub-id-type="pmid">10964230</pub-id></citation></ref>
<ref id="B46">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rich</surname> <given-names>T. C. G.</given-names>
</name>
<name>
<surname>Green</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Houston</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Lep&#x161;&#xed;</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Ludwig</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Pellicer</surname> <given-names>J.</given-names>
</name>
</person-group> (<year>2014</year>). <article-title>British <italic>Sorbus</italic> (Rosaceae): six new species, two hybrids and a new subgenus</article-title>. <source>New J. Bot.</source> <volume>4</volume>, <fpage>2</fpage>&#x2013;<lpage>12</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1179/2042349714Y.0000000036</pub-id>
</citation></ref>
<ref id="B47">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rich</surname> <given-names>T. C. G.</given-names>
</name>
<name>
<surname>Houston</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Wray</surname> <given-names>N.</given-names>
</name>
<name>
<surname>King</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Brown</surname> <given-names>A. P.</given-names>
</name>
<name>
<surname>Fay</surname> <given-names>M. F.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>1049. <italic>SORBUS BRISTOLIENSIS</italic>: rosaceae</article-title>. <source>Curtis&#x2019;s Bot. Mag.</source> <volume>39</volume>, <fpage>737</fpage>&#x2013;<lpage>751</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/curt.12483</pub-id>
</citation></ref>
<ref id="B48">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Robertson</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Newton</surname> <given-names>A. C.</given-names>
</name>
<name>
<surname>Ennos</surname> <given-names>R. A.</given-names>
</name>
</person-group> (<year>2004</year>). <article-title>Multiple hybrid origins, genetic diversity and population genetic structure of two endemic <italic>Sorbus</italic> taxa on the Isle of Arran, Scotland</article-title>. <source>Mol. Ecol.</source> <volume>13</volume>, <fpage>123</fpage>&#x2013;<lpage>134</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1046/j.1365-294X.2003.02025.x</pub-id>, PMID: <pub-id pub-id-type="pmid">14653794</pub-id></citation></ref>
<ref id="B49">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Robertson</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Rich</surname> <given-names>T. C. G.</given-names>
</name>
<name>
<surname>Allen</surname> <given-names>A. M.</given-names>
</name>
<name>
<surname>Houston</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Roberts</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Bridle</surname> <given-names>J. R.</given-names>
</name>
<etal/>
</person-group>. (<year>2010</year>). <article-title>Hybridization and polyploidy as drivers of continuing evolution and speciation in <italic>Sorbus</italic>
</article-title>. <source>Mol. Ecol.</source> <volume>19</volume>, <fpage>1675</fpage>&#x2013;<lpage>1690</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/j.1365-294X.2010.04585.x</pub-id>, PMID: <pub-id pub-id-type="pmid">20345692</pub-id></citation></ref>
<ref id="B50">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rozas</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Ferrer-Mata</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Sanchez-DelBarrio</surname> <given-names>J. C.</given-names>
</name>
<name>
<surname>Guirao-Rico</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Librado</surname> <given-names>P.</given-names>
</name>
<name>
<surname>Ramos-Onsins</surname> <given-names>S. E.</given-names>
</name>
<etal/>
</person-group>. (<year>2017</year>). <article-title>DnaSP 6: DNA sequence polymorphism analysis of large data sets</article-title>. <source>Mol. Biol. Evol.</source> <volume>34</volume>, <fpage>3299</fpage>&#x2013;<lpage>3302</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/molbev/msx248</pub-id>, PMID: <pub-id pub-id-type="pmid">29029172</pub-id></citation></ref>
<ref id="B51">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rushforth</surname> <given-names>K.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>The Whitebeam problem, and a solution</article-title>. <source>Phytologia</source> <volume>100</volume>, <fpage>222</fpage>&#x2013;<lpage>247</lpage>.</citation></ref>
<ref id="B52">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sennikov</surname> <given-names>A. N.</given-names>
</name>
<name>
<surname>Kurtto</surname> <given-names>A.</given-names>
</name>
</person-group> (<year>2017</year>). <article-title>A phylogenetic checklist of <italic>Sorbus</italic> s.l. (Rosaceae) in Europe</article-title>. <source>Memo. Soc pro Fauna Flora Fenn.</source> <volume>93</volume>, <fpage>1</fpage>&#x2013;<lpage>78</lpage>.</citation></ref>
<ref id="B53">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Shi</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Chen</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Jiang</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Wu</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Huang</surname> <given-names>L.</given-names>
</name>
<etal/>
</person-group>. (<year>2019</year>). <article-title>CPGAVAS2, an integrated plastome sequence annotator and analyzer</article-title>. <source>Nucleic Acids Res.</source> <volume>47</volume>, <fpage>W65</fpage>&#x2013;<lpage>W73</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/nar/gkz345</pub-id>, PMID: <pub-id pub-id-type="pmid">31066451</pub-id></citation></ref>
<ref id="B54">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Shi</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Chen</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Jiang</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Wu</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Xin</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Zeng</surname> <given-names>W.</given-names>
</name>
</person-group> (<year>2024</year>). <article-title>Assembly and comparative analysis of the complete mitogenome of <italic>Rubus chingii</italic> var. <italic>suavissimus</italic>, an exceptional berry plant possessing sweet leaves</article-title>. <source>Front. Plant Sci.</source> <volume>15</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fpls.2024.1504687</pub-id>, PMID: <pub-id pub-id-type="pmid">39764230</pub-id></citation></ref>
<ref id="B55">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Skippington</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Barkman</surname> <given-names>T. J.</given-names>
</name>
<name>
<surname>Rice</surname> <given-names>D. W.</given-names>
</name>
<name>
<surname>Palmer</surname> <given-names>J. D.</given-names>
</name>
</person-group> (<year>2015</year>). <article-title>Miniaturized mitogenome of the parasitic plant <italic>Viscum scurruloideum</italic> is extremely divergent and dynamic and has lost all nad genes</article-title>. <source>Proc. Natl. Acad. Sci.</source> <volume>112</volume>, <fpage>E3515</fpage>&#x2013;<lpage>E3524</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1073/pnas.1504491112</pub-id>, PMID: <pub-id pub-id-type="pmid">26100885</pub-id></citation></ref>
<ref id="B56">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Somlyay</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Sennikov</surname> <given-names>A. N.</given-names>
</name>
</person-group> (<year>2015</year>). <article-title>Taxonomic Interpretation and Typification of <italic>Sorbus pannonica</italic> (Rosaceae), a Presumed Intermediate between S. aria and S. graeca from Hungary</article-title>. <source>Ann. Bot. Fenn.</source> <volume>52</volume>, <fpage>274</fpage>&#x2013;<lpage>287</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.5735/085.052.0322</pub-id>
</citation></ref>
<ref id="B57">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sun</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Chen</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Liu</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Liu</surname> <given-names>B.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>J.</given-names>
</name>
<etal/>
</person-group>. (<year>2022</year>). <article-title>Rearrangement and domestication as drivers of Rosaceae mitogenome plasticity</article-title>. <source>BMC Biol.</source> <volume>20</volume>, <fpage>1</fpage>&#x2013;<lpage>19</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1186/s12915-022-01383-3</pub-id>, PMID: <pub-id pub-id-type="pmid">35986276</pub-id></citation></ref>
<ref id="B58">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sun</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Han</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Shen</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Liu</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Fan</surname> <given-names>W.</given-names>
</name>
<etal/>
</person-group>. (<year>2024</year>). <article-title>Comprehensive analysis of the Lycopodium japonicum mitogenome reveals abundant tRNA genes and cis-spliced introns in Lycopodiaceae species</article-title>. <source>Front. Plant Sci.</source> <volume>15</volume>, <elocation-id>1446015</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fpls.2024.1446015</pub-id>, PMID: <pub-id pub-id-type="pmid">39228832</pub-id></citation></ref>
<ref id="B59">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wang</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Qiu</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Sun</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Han</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Yang</surname> <given-names>Y.</given-names>
</name>
<etal/>
</person-group>. (<year>2025</year>). <article-title>Characterization and comparative analysis of the first mitochondrial genome of <italic>Michelia</italic> (Magnoliaceae)</article-title>. <source>Genomics Commun.</source> <volume>2</volume>, <elocation-id>e001</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.48130/gcomm-0025-0001</pub-id>
</citation></ref>
<ref id="B60">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wang</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Qin</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Shen</surname> <given-names>X.</given-names>
</name>
<name>
<surname>You</surname> <given-names>C.</given-names>
</name>
</person-group> (<year>2024</year>). <article-title>Morphological Structure Identification, Comparative Mitochondrial Genomics and Population Genetic Analysis toward Exploring Interspecific Variations and Phylogenetic Implications of <italic>Malus baccata</italic> &#x2018;ZA&#x2019; and Other Species</article-title>. <source>Biomolecules</source> <volume>14</volume>, <elocation-id>912</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/biom14080912</pub-id>, PMID: <pub-id pub-id-type="pmid">39199300</pub-id></citation></ref>
<ref id="B61">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wang</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Wu</surname> <given-names>Y.-W.</given-names>
</name>
<name>
<surname>Shih</surname> <given-names>A. C.-C.</given-names>
</name>
<name>
<surname>Wu</surname> <given-names>C.-S.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>Y.-N.</given-names>
</name>
<name>
<surname>Chaw</surname> <given-names>S.-M.</given-names>
</name>
</person-group> (<year>2007</year>). <article-title>Transfer of chloroplast genomic DNA to mitochondrial genome occurred at least 300 MYA</article-title>. <source>Mol. Biol. Evol.</source> <volume>24</volume>, <fpage>2040</fpage>&#x2013;<lpage>2048</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/molbev/msm133</pub-id>, PMID: <pub-id pub-id-type="pmid">17609537</pub-id></citation></ref>
<ref id="B62">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wee</surname> <given-names>C.-C.</given-names>
</name>
<name>
<surname>Nor Muhammad</surname> <given-names>N. A.</given-names>
</name>
<name>
<surname>Subbiah</surname> <given-names>V. K.</given-names>
</name>
<name>
<surname>Arita</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Nakamura</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Goh</surname> <given-names>H.-H.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>Plastomes of <italic>Garcinia mangostana</italic> L. and comparative analysis with other <italic>Garcinia</italic> species</article-title>. <source>Plants</source> <volume>12</volume>, <elocation-id>930</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/plants12040930</pub-id>, PMID: <pub-id pub-id-type="pmid">36840278</pub-id></citation></ref>
<ref id="B63">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wick</surname> <given-names>R. R.</given-names>
</name>
<name>
<surname>Schultz</surname> <given-names>M. B.</given-names>
</name>
<name>
<surname>Zobel</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Holt</surname> <given-names>K. E.</given-names>
</name>
</person-group> (<year>2015</year>). <article-title>Bandage: Interactive visualization of <italic>de novo</italic> genome assemblies</article-title>. <source>Bioinformatics</source> <volume>31</volume>, <fpage>3350</fpage>&#x2013;<lpage>3352</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/bioinformatics/btv383</pub-id>, PMID: <pub-id pub-id-type="pmid">26099265</pub-id></citation></ref>
<ref id="B64">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wu</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Liao</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Tembrock</surname> <given-names>L. R.</given-names>
</name>
<name>
<surname>Broz</surname> <given-names>A.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>Genomic architectural variation of plant mitochondria&#x2014;A review of multichromosomal structuring</article-title>. <source>J. Syst. Evol.</source> <volume>60</volume>, <fpage>160</fpage>&#x2013;<lpage>168</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/jse.12655</pub-id>
</citation></ref>
<ref id="B65">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wu</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Sloan</surname> <given-names>D. B.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>Recombination and intraspecific polymorphism for the presence and absence of entire chromosomes in mitochondrial genomes</article-title>. <source>Heredity (Edinb).</source> <volume>122</volume>, <fpage>647</fpage>&#x2013;<lpage>659</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/s41437-018-0153-3</pub-id>, PMID: <pub-id pub-id-type="pmid">30356223</pub-id></citation></ref>
<ref id="B66">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wu</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Yu</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Tang</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Yang</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Fu</surname> <given-names>Q.</given-names>
</name>
<name>
<surname>Zheng</surname> <given-names>Y.</given-names>
</name>
<etal/>
</person-group>. (<year>2024</year>). <article-title>Morphological and molecular evidence for natural hybridization between <italic>Sorbus pohuashanensis</italic> and <italic>S. discolor</italic> (Rosaceae)</article-title>. <source>J. For. Res.</source> <volume>35</volume>, <fpage>1</fpage>&#x2013;<lpage>13</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s11676-023-01659-6</pub-id>
</citation></ref>
<ref id="B67">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wynn</surname> <given-names>E. L.</given-names>
</name>
<name>
<surname>Christensen</surname> <given-names>A. C.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>Repeats of unusual size in plant mitochondrial genomes: identification, incidence and evolution</article-title>. <source>G3 Genes|Genomes|Genetics</source> <volume>9</volume>, <fpage>549</fpage>&#x2013;<lpage>559</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1534/g3.118.200948</pub-id>, PMID: <pub-id pub-id-type="pmid">30563833</pub-id></citation></ref>
<ref id="B68">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Yang</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Ni</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Lin</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Yang</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Chen</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Nijiati</surname> <given-names>N.</given-names>
</name>
<etal/>
</person-group>. (<year>2022</year>). <article-title>
<italic>De novo</italic> assembly of the complete mitochondrial genome of sweet potato (<italic>Ipomoea batatas</italic> [L.] Lam) revealed the existence of homologous conformations generated by the repeat-mediated recombination</article-title>. <source>BMC Plant Biol.</source> <volume>22</volume>, <fpage>1</fpage>&#x2013;<lpage>12</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1186/s12870-022-03665-y</pub-id>, PMID: <pub-id pub-id-type="pmid">35681138</pub-id></citation></ref>
<ref id="B69">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>You</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Cui</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Zang</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Su</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Que</surname> <given-names>Y</given-names>
</name>
</person-group>. (<year>2022</year>). <article-title>Assembly of the Complete Mitochondrial Genome of Gelsemium elegans Revealed the Existence of Homologous Conformations Generated by a Repeat Mediated Recombination</article-title>. <source>Int. J. Mol. Sci</source>. <volume>24</volume>, <elocation-id>527</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/ijms24010527</pub-id>, PMID: <pub-id pub-id-type="pmid">36613970</pub-id></citation></ref>
<ref id="B70">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Y&#xfc;</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Lu</surname> <given-names>L.</given-names>
</name>
</person-group> (<year>1974</year>). &#x201c;<article-title>
<italic>Sorbus</italic> L</article-title>,&#x201d; in <source>Flora Reipublicae Popularis Sinicae</source>, vol. <volume>36</volume> . Ed. <person-group person-group-type="editor">
<name>
<surname>Y&#xfc;</surname> <given-names>T. T.</given-names>
</name>
</person-group> (<publisher-name>Science Press</publisher-name>, <publisher-loc>Beijing</publisher-loc>), <fpage>283</fpage>&#x2013;<lpage>344</lpage>.</citation></ref>
<ref id="B71">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Zhai</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Zhao</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Fu</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Huang</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Jiang</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>M.</given-names>
</name>
<etal/>
</person-group>. (<year>2025</year>). <article-title>
<italic>De novo</italic> assembly and comparative analysis of cherry (<italic>Prunus</italic> subgenus <italic>Cerasus</italic>) mitogenomes</article-title>. <source>Front. Plant Sci.</source> <volume>16</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fpls.2025.1568698</pub-id>, PMID: <pub-id pub-id-type="pmid">40196431</pub-id></citation></ref>
<ref id="B72">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Zhang</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Gao</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Jakovli&#x107;</surname> <given-names>I.</given-names>
</name>
<name>
<surname>Zou</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>W. X.</given-names>
</name>
<etal/>
</person-group>. (<year>2020</year>). <article-title>PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies</article-title>. <source>Mol. Ecol. Resour.</source> <volume>20</volume>, <fpage>348</fpage>&#x2013;<lpage>355</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/1755-0998.13096</pub-id>, PMID: <pub-id pub-id-type="pmid">31599058</pub-id></citation></ref>
<ref id="B73">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Zhang</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Morales-Briones</surname> <given-names>D. F.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Huang</surname> <given-names>C. H.</given-names>
</name>
<etal/>
</person-group>. (<year>2023</year>). <article-title>Phylogenomics insights into gene evolution, rapid species diversification, and morphological innovation of the apple tribe (Maleae, Rosaceae)</article-title>. <source>New Phytol.</source> <volume>240</volume>, <fpage>2102</fpage>&#x2013;<lpage>2120</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/nph.19175</pub-id>, PMID: <pub-id pub-id-type="pmid">37537712</pub-id></citation></ref>
<ref id="B74">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Zhang</surname> <given-names>Q.-J.</given-names>
</name>
<name>
<surname>Wu</surname> <given-names>X.-Y.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>X.</given-names>
</name>
<name>
<surname>Yang</surname> <given-names>A.-S.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>X.-Y.</given-names>
</name>
<name>
<surname>Zhao</surname> <given-names>W.-M.</given-names>
</name>
<etal/>
</person-group>. (<year>2024</year>). <article-title>Evolutionary history and population dynamics of a rare and endangered medicinal plant <italic>Bergenia scopulosa</italic> (Saxifragaceae): Evidences from chloroplast genomes and ecological niche analysis</article-title>. <source>Glob. Ecol. Conserv.</source> <volume>54</volume>, <elocation-id>e03097</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.gecco.2024.e03097</pub-id>
</citation></ref>
<ref id="B75">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Zhang</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Yan</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Li</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Jiang</surname> <given-names>Z.</given-names>
</name>
<name>
<surname>Xiong</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>Y.</given-names>
</name>
<etal/>
</person-group>. (<year>2025</year>). <article-title>Assembly and comparative analysis of the complete mitochondrial genome of red raspberry (<italic>Rubus idaeus</italic> L.) revealing repeat-mediated recombination and gene transfer</article-title>. <source>BMC Plant Biol.</source> <volume>25</volume>, <fpage>85</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1186/s12870-024-05969-7</pub-id>, PMID: <pub-id pub-id-type="pmid">39838290</pub-id></citation></ref>
<ref id="B76">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Zheng</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Zhang</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2007</year>). <article-title>A cladistic and phenetic analysis of the infrageneric relationships of <italic>Sorbus</italic> s.l. (Maloideae, Rosaceae) based on the morphological characters</article-title>. <source>Acta Hortic. Sinnica</source> <volume>34</volume>, <fpage>723</fpage>&#x2013;<lpage>728</lpage>.</citation></ref>
<ref id="B77">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Zhou</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Armijos</surname> <given-names>C. E.</given-names>
</name>
<name>
<surname>Lee</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Lu</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Wang</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Ruhlman</surname> <given-names>T. A.</given-names>
</name>
<etal/>
</person-group>. (<year>2023</year>). <article-title>Plastid genome assembly using long-read data</article-title>. <source>Mol. Ecol. Resour.</source> <volume>23</volume>, <fpage>1442</fpage>&#x2013;<lpage>1457</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/1755-0998.13787</pub-id>, PMID: <pub-id pub-id-type="pmid">36939021</pub-id></citation></ref>
</ref-list>
</back>
</article>