Abstract
Within the genus Bellevalia (Asparagaceae: Scilloideae: Hyacintheae), flower shape and colour, the ratio of leaf to scape length, and the orientation of the pedicels were traditionally used to delimit sections, subsections and species. We conducted a phylogenetic analysis of representative species from sections Bellevalia, Conicae, Nutantes and Oxydontae, representing four out of the six sections that are currently recognised, using the four chloroplast regions rbcL, matK, trnL intron, and trnL-F spacer. Our results indicate that the sections are not monophyletic as currently circumscribed. Our analyses retrieve two major, well-supported clades. The first clade (sect. Conicae) includes only species with ciliate leaves and green-veined perianth lobes, and the second clade (sect. Bellevalia) includes the species lacking these characteristics. Within the second clade, the species with yellow anthers are separated from those with violet anthers. Our molecular analysis does not support the traditional subdivisions of the genus but establishes the value of cilia on the leaf margin, green veins in the perianths lobes, and the colour of the anthers as indicators of relationships among the species. We recommend that the current four sections and six subsections of Bellevalia be reduced to two sections without subsections and we provide an updated and corrected nomenclature for these sections and subsections, designating lectotypes where necessary.
References
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<p>Karabacak, O., Yıldırım, H. & Martin, E. (2015) <em>Bellevalia koyuncui sp. nova</em> (Asparagaceae): a new species from South Eastern Anatolia, Turkey. <em>Phytotaxa</em> 203 (1): 81–84. https://doi.org/10.11646/phytotaxa.203.1.9</p>
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<p>Angiosperm phylogeny Group [APGIV] (2016) An update of the Angiosperm phylogeny Group classification for the orders and families of flowering plants: APG IV. <em>Botanical Journal of the Linnean society </em>181: 1–20. https://doi.org/10.1111/boj.12385</p>
<p>Austuti, G., Brullo, S., Domina, G., El Mokni, R., Giordani, T. & Peruzzi, L. (2017) Phylogenetic relationships among tetraploid species of <em>Bellevalia</em> (Asparagaceae) endemic to south-central Mediterranean. <em>Plant Biosystems </em>151 (6): 1120–1128. https://doi.org/10.1080/11263504.2017.1320308</p>
<p>Bareka, P., Turland, N.J. & Kamari, G. (2015) <em>Bellevalia juliana</em> (Asparagaceae), a new hexaploid species from E Kriti (Greece). <em>Plant Biosystems</em> 149 (4): 703–709. https://doi.org/10.1080/11263504.2015.1057258</p>
<p>Boissier, E. (1854) <em>Diagnoses Plantarum Orientalium novarum</em>. B. Hermann, Lipsiae, 607 pp.</p>
<p>Boissier, E. (1884) <em>Diagnosis Plantarum Orientales novarum</em>. <em>Bulletin de l‘Herbier Boissier</em>.ser. 1, 5: 300–308. [Geneva]</p>
<p>Bornmüller, J.F.N. (1908) Beihefte zum Botanischen Centralblatt. Abt. 2, <em>Systematik, Pflanzengeographie, angewandte Botanik</em> 24 (2): 108. Kassel. Dresden.</p>
<p>Borzatti von Loewenstern, A., Giordani, T., Astuti, G., Andreucci, A. & Peruzzi, L. (2013) Phylogenetic relationships of Italian Bellevalia species (Asparagaceae), inferred from morphology, karyology and molecular systematics. <em>Plant Biosystems</em> 147: 776–787. https://doi.org/10.1080/11263504.2013.829884</p>
<p> Cowley, J., Özhatay, N. & Mathew, B. (1994) New species of Alliaceae & Hyacinthaceae from Turkey. <em>Kew Bulletin</em> 49: 481–489. https://doi.org/10.2307/4114472</p>
<p>Czernik (1923) <em>Hyacinthus litwinowii</em>., <em>Botanicheskie Materialy Gerbariya Glavnogo Botanicheskogo Sada R.S.F.S.R</em>. Petrograd 4: 148.</p>
<p>Doyle, J.J. & Doyle, J.L. (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. <em>Phytochemical Bulletin</em> 19: 11–15.</p>
<p>Dunning, L.T. & Savolainen, V. (2010) Broad-scale amplification of <em>mat</em>K for DNA barcoding plants, a technical note. <em>Botanical Journal of the Linnean Society</em> 164: 1–9. https://doi.org/10.1111/j.1095-8339.2010.01071.x</p>
<p>Engler, A. (1887) Liliaceae. <em>In:</em> <em>Die NaturlichenPflanzenfamilien, </em>vol. 2 (5). Leipzig, pp. 10–91.</p>
<p>Fay, M.F., Bayer, C., Alverson, W.S., de Bruijn, A.Y. & Chase, M.W. (1998) Plastid <em>rbc</em>L sequence data indicate a close affinity between <em>Diegodendron</em> and <em>Bixa</em>. <em>Taxon</em> 47: 43–50. https://doi.org/10.2307/1224017</p>
<p>Feinbrun, N. (1940) A monographic study of <em>Bellevalia </em>Lapeyr. <em>Palestine journal of botany</em> Jerusalem, ser 1: 336–340</p>
<p>Feinbrun, N. (1986) <em>Flora of Palestine. </em>The Israel academy of science & Humanity. <em>Jerusalem</em> 4: 84–104.</p>
<p>Freitag, H. & Wendelbo, P. (1970) The genus <em>Bellevalia</em> in Afghanistan. Studies in the Flora of Afghanistan. <em>Israel Journal of Botany</em> 19: 220–224.</p>
<p>Govaerts, R. (2018) <em>World checklist of Asparagaceae</em>. Available from: http://wcsp.science.kew.org (accessed 11 November 2021)</p>
<p>Groossheim, A.A. (1928) <em>Flora Kavkaza</em>, vol. 1. Izdatel’stvo Akademii Nauk SSSR, Moskva & Leningrad, 234 pp.</p>
<p>Gürdal, B., Özhatay, N., Koçyiğit, M. & Kaya, E. (2014) Two new species for Turkish geophyte flora in the genus <em>Bellevalia</em>. <em>In</em>: Kaya, E. (Ed.) Türkiye Geofitleri 3, Atatürk Bahçe Kültürleri Merkez Araştırma Enstitüsü, Yalova, pp. 525–531.</p>
<p>Gussone, G. (1841) <em>Muscari neglectum</em>. <em>Annali Civili del Regno delle due Sicilie</em>. Naples 26: 48–49.</p>
<p>Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analyses program for Windows 95/98/NT. <em>Nucleic Acids Symposium Series</em> 41: 95–98.</p>
<p>Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. <em>Bioinformatics</em> 17 (8): 754–755. https://doi.org/10.1093/bioinformatics/17.8.754</p>
<p>Jafari, A. & Maassoumi, A.A. (2008) A new species of <em>Bellevalia</em> (Liliaceae/ Hyacinthaceae) from Iran. <em>Edinburgh journal of Botany</em> 65 (3): 469–473. https://doi.org/10.1017/S0960428608005027</p>
<p>Karabacak, O., Yıldırım, H. & Martin, E. (2014) <em>Bellevalia pseudolongipes</em> <em>sp. nov</em>. (Asparagaceae): a new species from southeastern Anatolia, Turkey. <em>Phytotaxa</em> 188 (4): 209–217. https://doi.org/10.11646/phytotaxa.188.4.3</p>
<p>Karabacak, O., Yıldırım, H. & Martin, E. (2015) <em>Bellevalia koyuncui sp. nova</em> (Asparagaceae): a new species from South Eastern Anatolia, Turkey. <em>Phytotaxa</em> 203 (1): 81–84. https://doi.org/10.11646/phytotaxa.203.1.9</p>
<p>Kunth, C.S. (1843) <em>Bellevalia glauca</em>. <em>Enumeratio Plantarum</em> 4: 309.</p>
<p>Lapeyrouse, P.P. (1808) <em>Bellevalia</em> Lapeyr. Nouveau genre de plante de la famille des Liliacées<em>. Journal de Physique, de Chimie, d’Historie Naturelle et des Arts</em> 67 (12): 425–427.</p>
<p>Linneì, C. (1771) <em>Mantissa plantarum altera</em>. Holmiae : Impensis Direct. Laurentii Salvii, 224 pp.</p>
<p>Losinskaja, A.S.L. (1935) <em>Bellevalia</em> Lapeyr. <em>In</em>: Komarov, V.L. (Ed.) <em>Flora of USSR</em> 4. Akademiya Nauk SSSR., Leningrad, pp. 303–311.</p>
<p>Manning, C., Forest, F., Devey, D.S., Fay, M. & Goldblatt, P. (2009) A molecular phylogeny and a revised classification of Ornithogaloideae (Hyacinthaceae) based on an analysis of four plastid DNA regions. <em>Taxon</em> 58 (1): 1–107. https://doi.org/10.1002/tax.581011</p>
<p>Müller, K. (2005) SeqState - primer design and sequence statistics for phylogenetic DNA data sets. <em>Applied Bioinformatics</em> 4: 65–69. https://doi.org/10.2165/00822942-200504010-00008</p>
<p>Nylander, J.A.A. (2004) <em>MrModeltest, version 2. Program distributed by the author</em>. Evolutionary Biology Centre, Uppsala University.</p>
<p>Pallas, P.S. (1927) <em>Hyacinthus sarmaticus</em> Pall. ex Miscz. <em>Trudy Kubanskii Sel’skokhoz. Inst</em>. 5: 155.</p>
<p>Parlatore, F. (1854) <em>Bellevalia webbiana</em>. Nouv. Gen. Sp. Monocot, 19 pp.</p>
<p>Persson, K. & Wendelbo, P. (1979) The taxonomic position of <em>Bellevalia tabriziana</em> (Liliaceae). <em>Botaniska Notiser</em> 132: 197–200.</p>
<p>Pfosser, M. & Speta, F. (1999) Phylogenetic of Hyacinthaceae based on plastid DNA sequences. <em>Annals of the Missouri Botanical Garden</em> 86 (4): 852–875. https://doi.org/10.2307/2666172</p>
<p>Pinar, S.M. & Eroğlu, H. & Fidan, M. (2016) <em>Bellevalia behcetii sp. nov.</em> (Asparagaceae): a new species from South Eastern Anatolia, Turkey. <em>Phytotaxa</em> 270 (2): 127–136. https://doi.org/10.11646/phytotaxa.270.2.5</p>
<p>Rambaut, A. & Drummond, A. (2009) Tracer. version 1.5. [http://tree.bio.ed.ac.uk/software/ tracer]</p>
<p>Reeves, G., Chase, M.W. Goldblatt, P., Rudall, P., Fay, M.F., Cox, A.V., Lejeune, B. & Souza-Chies, T. (2001) Molecular systematic of Iridaceae: Evidence from four plastid DNA regions. <em>American Journal of Botany</em> 88 (11): 2074–2087. https://doi.org/10.2307/3558433</p>
<p>Roemer, J.J., Schultes, J.A. & Schultes, J.H. (1830) <em>Systema Vegetabilium </em>ed. 15 bis 7. pp. 586.</p>
<p>Simmons, M.P. & Ochoterena, H. (2000) Gaps as characters in sequence-based phylogenetic analyses. <em>Systematic Biology</em> 49: 369–381. https://doi.org/10.1093/sysbio/49.2.369</p>
<p>Speta, F. (1998) Hyacinthaceae. <em>In</em>: Kubitzki, K. (Ed.) <em>The families and genera of vascular plants. III. Flowering plants. Monocotyledons. (except Orchidaceae</em>). Springer. Berlin, 261–285. https://doi.org/10.1007/978-3-662-03533-7_35</p>
<p>Taberlet, P., Gielly, L., Pautou, G. & Bouvet, J. (1991) Universal primers for amplification of three non-coding regions of chloroplast DNA. <em>Plant Molecular Biology</em> 17: 1105–1109. https://doi.org/10.1007/BF00037152</p>
<p>Thiers, B. (2016 [continuously updated])<em> Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium</em>. Available from: http://sweetgum.nybg.org/ih/ (accessed 7 May 2016)</p>
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