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Phylogeny and biogeography of the genus Hesperis (Brassicaceae, tribe Hesperideae) inferred from nuclear ribosomal DNA sequence data

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Abstract

Despite its complicated taxonomy, the genus Hesperis has not yet been subjected to any detailed molecular phylogenetic study and little is known about its monophyly, origin and biogeographical history, as well as the evolution of morphological characters. Here, we present for the first time comprehensive molecular analyses (nuclear ribosomal ITS sequences) of approximately 40 Hesperis species which represent the full range of morphological variation and the entire geographic distribution area of the genus. Based on our results, monophyly of the genus Hesperis has been proved. Moreover, our phylogenetic analysis shows that almost all traditionally defined sections are not monophyletic. NeighborNet network analysis was performed in order to visualize conflicting phylogenetic signals. The split-graph corresponds to the major clades represented in the consensus tree and revealed a hybridization signal in the evolutionary history of only a single Hesperis species. Divergence time estimations indicate that the origin of Hesperis (7.66–19.9 Mya) coincides with the expansion of grasslands and the closure of the proto-Mediterranean Sea in the middle Miocene. Therefore, diversification within Hesperis was affected by global climate changes, substantial tectonic rearrangements and the expansion of open vegetation systems in the Miocene, all of which had a great impact on the speciation history of the Irano-Turanian flora and fauna. Furthermore, two new species from West of Iran are described and illustrated here. They are readily distinguished from their closely related species by morphological characters such as stem height, leaf shape, pedicel, sepal and fruit indumentum, petal size, stigma shape and seed characters.

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Acknowledgements

This project was carried out at Hacettepe University by the first author (AEF) who wishes to express her special thanks to Hacettepe University for providing financial support and lab facilities. The authors thank the curators of the following institutions: the Research Institute of Forests and Rangelands (TARI), the Herbarium of Shiraz University (HSHU), the Hacettepe University Herbarium (HUB), the Konya Herbarium (KNYA), the Royal Botanic Gardens, Kew (K) & the Herbarium Russian Academy of Sciences- V. L. Komarov Botanical institute (LE) for providing herbarium materials. We also appreciate Dr. Sara Manafzadeh´s advice in selecting the appropriate method in biogeographical studies and Dr. Ihsan Al-Shehbaz for preparing Caucasian plant materials for phylogenetic study and his valuable recommendations. Special thanks go to Dr. Dmitry A. German for his valuable comments on the taxonomic part of the paper, and also special thanks to Prof. Dr. Ahmad Reza Khosravi for his great cooperation in the study of Sisymbrium leucocladum specimens in Shiraz University Herbarium. Furthermore, we thank Robabeh Farahdust, the artist from the Central Herbarium of Iran for preparing the sketches of the new species. Finally, we would like to express our deep gratitude to Lucille Schmieding for her generous support in language edition.

Funding

Hacettepe University provided financial support and laboratory facilities; Lorestan University financially supported the SEM studies.

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Correspondence to Hamed Khodayari.

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Online Resource 1. GenBank accession numbers for the sequences (ITS + trnL-F region) used in phylogenetic study.

Online Resource 2. DNA sequence alignments of ITS dataset of Hesperis specimens for the current study in NEXUS format.

Online Resource 3. DNA sequence alignments of trnL-F dataset of Hesperis specimens for the current study in NEXUS format.

Online Resource 4. List of the character states used in the Ancestral Character State Reconstruction analysis.

Online Resource 5. Statistical output of ancestral area reconstruction analysis based on (A) Bayesian Binary MCMC (BBM) and (B) BioGeoBears (BayAreaLike) of the genus Hesperis.

Online Resource 6. Bayesian 50% majority rule consensus tree inferred from the trnL-F dataset.

Online Resource 7. Hesperis ilamica. a Plant, b flower, c petal (frontal view), d long stamen, e short stamen, f outer sepal, g inner sepal, h fruit, i fruit apex (drawing prepared from TARI, No. 102982).

Online Resource 8. Hesperis bakhtiarica. a Plant, b inlorescence, c flower, d outer sepal, e inner sepal, f long stamen, g short stamen, h petal (frontal view), i fruit apex (drawing prepared from TARI, No. 59927).

Online Resource 9. Hesperis ilamica, sp. nov. a Habitat, b–c, e flower, d stigma, f–g herbarium specimens including holotypus.

Online Resource 10. General view and habitat of Hesperis bakhtiarica, sp. nov. a–b, d Herbarium specimens including holotypus, c flower, e habitat.

Online Resource 11. The SEM micrographs of Iranian Hesperis new species. a–i Hesperis bakhtiarica: a fruit, b leaf, c–d petal (dorsal and ventral, respectively), e stem, f stigma, g pollen grain, h–i seed; j–r H. ilamica: j fruit, k leaf, l–m petal (dorsal and ventral, respectively), n stem, o stigma, p pollen grain, q–r seed.

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Eslami-Farouji, A., Khodayari, H., Assadi, M. et al. Phylogeny and biogeography of the genus Hesperis (Brassicaceae, tribe Hesperideae) inferred from nuclear ribosomal DNA sequence data. Plant Syst Evol 307, 17 (2021). https://doi.org/10.1007/s00606-020-01727-y

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