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Allium Schisticola, a New Species with Peculiar Morphological Characters and an Unexpected Phylogenetic Position

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Allium Schisticola, a New Species with Peculiar Morphological Characters and an Unexpected Phylogenetic Position Phytotaxa 450 (3): 246–256 ISSN 1179-3155 (print edition) https://www.mapress.com/j/pt/ PHYTOTAXA Copyright © 2020 Magnolia Press Article ISSN 1179-3163 (online edition) https://doi.org/10.11646/phytotaxa.450.3.1 Allium schisticola, a new species with peculiar morphological characters and an unexpected phylogenetic position ALIREZA DOLATYARI1,4, HAMID MOAZZENI2,5*, SAEIDE HOSSEINI2,6, FRANK R. BLATTNER3,7 & REINHARD M. FRITSCH3,8 1 Plant Bank, Iranian Biological Resource Center (IBRC), Research Complex of ACECR (Iranian Academic Center for Education, Culture & Research), Karaj, Iran. 2 FUMH herbarium, Department of Botany, Research Center for Plant Sciences, Ferdowsi University of Mashhad, P.O. Box 91779- 48974, Mashhad, Iran. 3 Leibniz-Institut für Pflanzengenetik und Kulturpflanzenforschung (IPK), Corrensstrasse 3, D-06466 Seeland OT Gatersleben, Germany. 4 [email protected]; https://orcid.org/0000-0003-2866-1073 5 [email protected]; https://orcid.org/0000-0002-2406-2666 6 [email protected]; http://orcid.org/0000-0001-9367-5530 7 [email protected]; https://orcid.org/0000-0001-7731-7741 8 [email protected]; https://orcid.org/0000-0001-8207-7877 *Author for correspondence Abstract Allium schisticola is described from West Azarbaijan (Iran) as a new species. It is closely related to A. sabalense and A. sahandicum in having a similar flower color but differs by leaf, filament, and tepal characters. We investigated the phylogenetic relationship of the new species based on sequences of the chloroplast trnL-trnF and nuclear ribosomal DNA internal transcribed spacer (ITS) regions in A. subg. Melanocrommyum compared with 109 accessions of this subgenus. Our results confirm its placement in A. subg. Melanocrommyum. However, the ITS tree showed that the new species should belong to A. sect. Melanocrommyum despite the fact that it represents many morphological characters of A. sect. Acanthoprason. Most peculiar for the new species are obtuse tepals broadest near the tip (vs. very narrowly lanceolate up to triangular tepals in A. sect. Acanthoprason), with adaxially inconspicuous median vein (vs. conspicuous median vein in sect. Acanthoprason). The new species is diploid with a chromosome number of 2n = 2x = 16. Karyotype features and meiotic chromosomes behavior are presented for the new taxon. A detailed morphological description, illustrations, and a distribution map of the new species are given. Keywords: Allium sabalense, Allium sahandicum, Iran, phylogeny, section Acanthoprason, subg. Melanocrommyum Introduction The genus Allium Linnaeus (1753: 294) (Amaryllidaceae) with currently accepted 977 species (Govaerts et al. 2019) is one of the largest genera among the monocots. It is mainly distributed on the Northern Hemisphere from the Mediterranean Basin to Central Asia but adapted to diverse habitats (Fritsch 2012, Fritsch et al. 2010, Govaerts et al. 2019, Li et al. 2010). Most Allium species are perennial herbal geophytes characterized by tunicated bulbs, narrow to broad basal leaves, umbellate or head-like inflorescences, flowers mainly in pink, white and yellow; loculicidal capsules, rhomboidal or spheroidal, angled black seeds, and a specific odor and taste due to the presence of cysteine sulphoxides (Li et al. 2010). The genus has been classified into 15 subgenera and 85 sections (Friesen et al. 2006, Fritsch et al. 2010, Huang et al. 2014, Khassanov et al. 2011, Khassanov et al. 2013). Allium subg. Melanocrommyum (Webb & Berthelot 1848: 347) Rouy (1910: 378), with about 160 accepted species is the second largest subgenus of Allium (Fritsch 2016). Its members are naturally distributed from the Canary Islands to northwestern India, and most species occur in Asia Minor, Southwest Asia, and Central Asia (Fritsch 2016). Iran, with about 83 species (of over 155 Allium species growing naturally in Iran), belongs to the main center of diversity of this subgenus. Many members of this subgenus 246 Accepted by Lorenzo Peruzzi: 18 Jun. 2020; published: 30 Jun. 2020 can easily be recognized by having broad and flat leaves, rigid and most often strictly upright scapes of varying length and large fasciculate to globular inflorescences composed of many moderately small to large and often star-like flowers (Fritsch et al. 2010, Fritsch & Abbasi 2013). Allium subg. Melanocrommyum comprises ten sections in Iran, among which A. sect. Acanthoprason Wendelbo (1969: 27) with 25 accepted species is the largest section, morphologically recognized by relatively short peduncles and tepals forming a rigid spine-like median vein after anthesis (Fritsch & Abbasi 2009). Local people eat fresh or cooked young leaves of several members of A. sect. Acanthoprason (generally called “valak”). During a botanical excursion to northwest Iran, we found an Allium species growing on slate slopes. These specimens should belong to A. subg. Melanocrommyum, but we were not able to identify them using the keys and descriptions given in local Floras (Wendelbo 1971; Fritsch & Abbasi 2013, and references therein). They are described here as a new species. TABLE 1. Voucher specimens, origin and GenBank accessions of taxa newly sequenced in the present study. All vouchers are kept in IBRC herbarium. IBRC No./ Taxon origin ITS trnL-trnF Herbarium no. IRAN. Zanjan: Zanjan towards Gilvan, 5 km Allium alamutense Razyfard, Zarre before Khanchaei road guard station, 2400 & R.M.Fritsch in Razyfard et al. P1010547/1616 - MT130434 m, 25 May 2014, Dolatyari, Ramezani & (2011: 353) Abdolahi IRAN. Ilam: Malekshahi, 5 km before Allium austroiranicum R.M.Fritsch Meimeh, 6 km on the road towards P1010320/1663 - MT130435 in Fritsch & Abbasi (2009: 26) Kabirkuh, 2200 m, 3 May 2014 Moazzeni, Dolatyari & Ramezani IRAN. Ilam: Ilam towards Eyvan, Chovar, Allium chrysantherum Boiss. & Reut. Reno forest park, 2 km from junction P1010382/1630 - MT130436 in Boissier (1882: 280). towards Park, 1500 m, 5 May 2014, Moazzeni, Dolatyari & Ramezani IRAN. Kermanshah: Eslam Abad-e Qarb Allium koelzii (Wendelbo 1966: 53) towards Mahidasht, Mersad strait, 1500 m, 6 P1010401/1607 - MT130437 Persson & Wendelbo (1979: 196) May 2014 Moazzeni, Dolatyari & Ramezani IRAN. West Azerbaijan: 61 km before Allium latifolium Jaub. & Spach Orumieh from Salmas, 1842 m, 6 June 2013, P1010022/1612 - MT130438 (1844: 2) Moazzeni, Dolatyari & Ramezani IRAN. Zanjan: Dandi towards Takht-e Allium mahneshanense Razyfard, Soleiman, NW mountain of lead and zinc Zarre & R.M.Fritsch in Razyfard et P1010506/1656 - MT130439 Anguran mine, 2850 m, 23 May 2014, al. (2011: 352) Dolatyari, Ramezani & Abdolahi IRAN. East Azerbaijan: S of Ahar, 3 km after Kaghalagh, Ghosh-e Dagh altitudes. Allium materculae Bordz. (1915: 73) P1009837/1834 - MT130440 2000 m, 2 June 2013, Moazzeni, Dolatyari & Ramezani Allium pseudohollandicum IRAN. Zanjan: Dandi towards Takab, 55 R.M.Fritsch in Fritsch & Abbasi km before Takab, 2500 m, 22 May 2014, P1010525/1638 - MT130441 (2013: 158) Dolatyari, Ramezani & Abdolahi IRAN. West Azerbaijan: 20 km from Khoy MT112908 Allium schisticola R.M. Fritsch, towards Qotor, 7 km towards Almalu- P1009984/3385 ‒ MT130443 Moazzeni & Dolatyari Hendowan villages, 1800 m, 5 June 2013, MT112917 Moazzeni, Dolatyari & Ramezani IRAN. Zanjan: Soltaniyeh, Arjin, N slope of Allium ubipetrense R.M.Fritsch in mountain W of sand mine, 2300 m, 22 May P1010476/1615 - MT130442 Fritsch & Abbasi (2009: 20) 2014, Dolatyari, Ramezani & Abdolahi A NEW SPECIES OF ALLIUM SCHISTICOLA Phytotaxa 450 (3) © 2020 Magnolia Press • 247 FIGURE 1. Allium schisticola A–C: Cultivated plant at anthesis. D–F: Plants in their natural habitat; D: Plant in the fruiting stage; E: Close-up of fruits; F: Bulb; G: Seeds (Millimeter paper as background). All photos by Hamid Moazzeni. 248 • Phytotaxa 450 (3) © 2020 Magnolia Press DOLATYARI ET AL. FIGURE 2. Close relatives of the new species. A–B: Allium cardiostemon, A: Habit, B: Inflorescences; C–D: A. sabalense, C: Habit, D: Close-up of a flower. E–G: A. sahandicum, E: Close-up of a flower, F–G: Habit. All photos by Hamid Moazzeni. Material and methods Source of material:—Living plants of the new Allium species and other members of A. subg. Melanocrommyum A NEW SPECIES OF ALLIUM SCHISTICOLA Phytotaxa 450 (3) © 2020 Magnolia Press • 249 were collected during fieldwork and were planted in the research farm of Iranian Biological Resource Center (IBRC). Several herbarium specimens were also prepared for further morphological investigation. All the specimens were deposited at IBRC herbarium. DNA Isolation, Amplification, and Sequencing:—Total genomic DNA was extracted from silica dried materials of 20 specimens (10 individuals of the new species and 10 species of A. subg. Melanocrommyum) using the 2× cetyltrimethylammonium bromide (CTAB) extraction protocol (Doyle & Doyle 1987). Voucher information and GenBank accession numbers for all newly included specimens are listed in Table 1. The nuclear ribosomal DNA internal transcribed spacer (ITS) and the plastid trnL-trnF regions were amplified and sequenced for ten individuals of the new species and ten endemic species from A. subg. Melanocrommyum for trnL-trnF (for more details see Table 1). For ITS, ITS-1 and ITS-2 were amplified separately using amplification primers (Blattner 1999; for ITS-1, the primers ITS-A and ITS-C, for ITS-2 the primers ITS-Dmel and ITS-B) following Gurushidze et al. (2008). The trnL gene together with the trnL–trnF intergenic spacer was amplified using the primers of Taberlet et al. (1991) as described in Gurushidze et al. (2010). Amplification products for both markers were purified using PEG (Joly et al. 2006). Sequencing was performed by BGI (Hong Kong) using the PCR primers. Sequences were trimmed, assembled, and aligned using Geneious version 6.1.2 (https://www.geneious.com). The newly generated sequences for ten accessions of the new species and ten closely related species were submitted to GenBank (accession numbers MT112908 to MT112917 for ITS and MT130434 to MT130443 for trnL-trnF, see Table 1). The new ITS sequences corresponding to the new species were added to the ITS matrix from Fritsch & Abbasi (2013: 198-202). For trnL-trnF, all sequences of A. subg. Melanocrommyum were obtained from GenBank.
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