b3528a49daac40a810a12cd20581e398 Heimer_2025_SB.pdf 6a5a8d7f2c0d5bbc77b6fddbfb496c3a932c09f7 Heimer_2025_SB.pdf 70d3162be58679ad7bc64eff2033c9172c4bb483a733598019506507f38200b2 Heimer_2025_SB.pdf Title: Hybridization and Polyploidy Shaped the Evolutionary History of a Complex of Cryptic Species in European Woodrushes (Luzula sect. Luzula) Subject: DOI: 10.1093/sysbio/syaf065 Systematic Biology, 0, 0, 25-9-2025. Abstract: Polyploidization has played a central role in the evolutionary history of most plant lineages, yet it poses significant challenges for phylogenetic inference, particularly in allopolyploid complexes with reticulate species relationships. Luzula sect. Luzula (Juncaceae) is a taxonomically intricate group characterized by widespread polyploidy, agmatoploidy, and high morphological uniformity. Focusing on the Eastern Alps, a key center of its diversity, we collected 1002 samples of nine species and applied an integrative framework combining ddRADseq, plastid sequencing, relative genome size estimation, and chromosome counting to disentangle its evolutionary history. We extended previously inferred phylogenetic relationships and assessed gene flow among diploids, establishing a baseline for investigating the origin of polyploids. By analyzing patterns of genotype frequencies and genetic affinities to diploids, we inferred the most likely parental species of polyploids and identified key hybridization events shaping the current taxonomic and karyotypic diversity within this group. Our results reveal weak genetic differentiation among some diploid lineages, likely reflecting gene flow and incomplete lineage sorting. We propose a common allopolyploid origin of two tetraploids, which subsequently gave rise to a third tetraploid and a hexaploid species through interploidy hybridization. Although the parental species of some polyploids remain obscure, our genomic data highlight polyploidy and hybridization as major drivers of speciation in this poorly understood lineage. This study underscores the value of integrative approaches in resolving reticulate plant phylogenies and advances our understanding of polyploid speciation. Keywords: Author: Heimer Valentin, Carnicero Pau, Carrizo GarcÃ�a Carolina , Hilpold Andreas, Dolenc Koce Jasna , Leal J. Luis, Li Mingai, Varotto Claudio, Schönswetter Peter, Frajman Božo, Särkinen Tiina Creator: OUP Producer: Acrobat Distiller 25.0 (Windows); modified using iTextSharp.LGPLv2.Core 3.7.4.0 CreationDate: Fri Nov 14 12:51:17 2025 CET ModDate: Mon Nov 24 11:36:08 2025 CET Custom Metadata: yes Metadata Stream: yes Tagged: yes UserProperties: no Suspects: no Form: none JavaScript: no Pages: 24 Encrypted: no Page size: 595.276 x 782.362 pts Page rot: 0 File size: 2901016 bytes Optimized: no PDF version: 1.4 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- EIDNBC+PalatinoLTStd-Roman Type 1C Custom yes yes yes 141 0 EIDNBD+dummy-space Type 1C WinAnsi yes yes no 142 0 EIDNBE+PalatinoLTStd-Italic Type 1C Custom yes yes yes 143 0 EIDNCF+PalatinoLTStd-Bold Type 1C Custom yes yes yes 6944 0 EIDNCG+PalatinoLTStd-BoldItalic Type 1C Custom yes yes yes 6945 0 EIECNE+MTSY Type 1C Custom yes yes yes 144 0 JZLQXJ+ArialMT TrueType WinAnsi yes yes no 1 0 EIEJGM+EURM10 Type 1C Custom yes yes yes 288 0 EIEMEM+RMTMI Type 1C Custom yes yes yes 396 0 EIHEME+Dingbats Type 1C Custom yes yes no 1549 0 Jhove (Rel. 1.28.0, 2023-05-18) Date: 2025-11-28 03:10:45 CET RepresentationInformation: Heimer_2025_SB.pdf ReportingModule: PDF-hul, Rel. 1.12.4 (2023-03-16) LastModified: 2025-11-27 04:02:04 CET Size: 2901016 Format: PDF Version: 1.4 Status: Well-Formed and valid SignatureMatches: PDF-hul MIMEtype: application/pdf Profile: Tagged PDF PDFMetadata: Objects: 7021 FreeObjects: 1 IncrementalUpdates: 0 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: false CenterWindow: false DisplayDocTitle: true NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: OneColumn PageMode: UseOutlines Language: English Outlines: Item: Title: Materials and Methods Destination: section.1.1.0 Item: Title: Results Destination: section.2.14.0 Item: Title: Discussion Destination: section.3.23.0 Item: Title: Conclusions Destination: section.4.31.0 Item: Title: Supplementary Material Destination: section.5.32.0 Item: Title: Funding Destination: section.6.33.0 Item: Title: Acknowledgments Destination: section.7.34.0 Item: Title: Author Contributions Destination: section.8.35.0 Item: Title: Data Availability Destination: section.9.36.0 Item: Title: Conflict of Interest Destination: section.10.37.0 Item: Title: References Destination: section.11.37.0 Info: Title: Hybridization and Polyploidy Shaped the Evolutionary History of a Complex of Cryptic Species in European Woodrushes (Luzula sect. Luzula) Author: Heimer Valentin, Carnicero Pau, Carrizo García Carolina , Hilpold Andreas, Dolenc Koce Jasna , Leal J. Luis, Li Mingai, Varotto Claudio, Schönswetter Peter, Frajman Božo, Särkinen Tiina Subject: DOI: 10.1093/sysbio/syaf065 Systematic Biology, 0, 0, 25-9-2025. Abstract: Polyploidization has played a central role in the evolutionary history of most plant lineages, yet it poses significant challenges for phylogenetic inference, particularly in allopolyploid complexes with reticulate species relationships. Luzula sect. Luzula (Juncaceae) is a taxonomically intricate group characterized by widespread polyploidy, agmatoploidy, and high morphological uniformity. Focusing on the Eastern Alps, a key center of its diversity, we collected 1002 samples of nine species and applied an integrative framework combining ddRADseq, plastid sequencing, relative genome size estimation, and chromosome counting to disentangle its evolutionary history. We extended previously inferred phylogenetic relationships and assessed gene flow among diploids, establishing a baseline for investigating the origin of polyploids. By analyzing patterns of genotype frequencies and genetic affinities to diploids, we inferred the most likely parental species of polyploids and identified key hybridization events shaping the current taxonomic and karyotypic diversity within this group. Our results reveal weak genetic differentiation among some diploid lineages, likely reflecting gene flow and incomplete lineage sorting. We propose a common allopolyploid origin of two tetraploids, which subsequently gave rise to a third tetraploid and a hexaploid species through interploidy hybridization. Although the parental species of some polyploids remain obscure, our genomic data highlight polyploidy and hybridization as major drivers of speciation in this poorly understood lineage. This study underscores the value of integrative approaches in resolving reticulate plant phylogenies and advances our understanding of polyploid speciation. 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Published by Oxford University Press on behalf of the Society of Systematic Biologists. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited. All Rights Reserved application/pdf doi:10.1093/sysbio/syaf065 © The Author(s) 2025. Published by Oxford University Press on behalf of the Society of Systematic Biologists. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited. Hybridization and Polyploidy Shaped the Evolutionary History of a Complex of Cryptic Species in European Woodrushes (Luzula sect. Luzula) DOI: 10.1093/sysbio/syaf065 Systematic Biology, 0, 0, 25-9-2025. Abstract: Polyploidization has played a central role in the evolutionary history of most plant lineages, yet it poses significant challenges for phylogenetic inference, particularly in allopolyploid complexes with reticulate species relationships. Luzula sect. Luzula (Juncaceae) is a taxonomically intricate group characterized by widespread polyploidy, agmatoploidy, and high morphological uniformity. Focusing on the Eastern Alps, a key center of its diversity, we collected 1002 samples of nine species and applied an integrative framework combining ddRADseq, plastid sequencing, relative genome size estimation, and chromosome counting to disentangle its evolutionary history. We extended previously inferred phylogenetic relationships and assessed gene flow among diploids, establishing a baseline for investigating the origin of polyploids. By analyzing patterns of genotype frequencies and genetic affinities to diploids, we inferred the most likely parental species of polyploids and identified key hybridization events shaping the current taxonomic and karyotypic diversity within this group. Our results reveal weak genetic differentiation among some diploid lineages, likely reflecting gene flow and incomplete lineage sorting. We propose a common allopolyploid origin of two tetraploids, which subsequently gave rise to a third tetraploid and a hexaploid species through interploidy hybridization. Although the parental species of some polyploids remain obscure, our genomic data highlight polyploidy and hybridization as major drivers of speciation in this poorly understood lineage. This study underscores the value of integrative approaches in resolving reticulate plant phylogenies and advances our understanding of polyploid speciation. Heimer Valentin Carnicero Pau Carrizo García Carolina Hilpold Andreas Dolenc Koce Jasna Leal J. Luis Li Mingai Varotto Claudio Schönswetter Peter Frajman Božo Särkinen Tiina Oxford University Press allopolyploidy ddRADseq Eastern Alps hybridization interploidy polyploid phylogenetics reticulate evolution Heimer Valentin, Carnicero Pau, Carrizo García Carolina , Hilpold Andreas, Dolenc Koce Jasna , Leal J. 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