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Tribe Eritrichieae (Boraginaceae s.str.) in West Asia: a molecular phylogenetic perspective

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Abstract

A selection of Boraginaceae genera was used to obtain a framework for the phylogenetic position of some tribes belong to subfamily Boraginoideae and genera within tribe Eritrichieae (Heterocaryum, Rochelia, Eritrichium, Lappula, Lepechiniella, and Asperugo) and related species. Our results were produced on the basis of nrDNA ITS and cpDNAtrnL-F sequences. The combined nrDNA ITS trnL-F data confirm four main clades of Boraginoideae comprising Echiochileae, Boragineae, Lithospermeae, and Cynoglosseae s. l. (including Eritrichieae, Cynoglosseae s. str., and Myosotideae). The tribe Eritrichieae itself at the current status is paraphyletic; some members, for example Asperugo procumbens, Lepechiniella inconspicua, Myosotidium hortensia, and Cryptantha flavoculata are placed out of the core tribe Eritrichieae. The genus Heterocaryum is monophyletic and allied with a subclade of genera Lappula, Lepechiniella, Eritrichium, and Rochelia. Rochelia is monophyletic, but Eritrichium and Lappula are non-monophyletic. Lepechiniella is nested among a group of Lappula species.

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Acknowledgments

We thank the herbariums TARI and FUMH for providing leaf materials.

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Correspondence to Maryam Khoshsokhan Mozaffar or Shahrokh Kazempour Osaloo.

Appendix

Appendix

Genbank accession numbers for nrDNA ITS and cpDNAtrnL-F sequence data, and voucher information with herbarium where specimen is deposited. The accession numbers for nrDNA ITS (reference or Location)/ and trnL-F (reference or location) are given in front of the species names. The dash symbol (–) shows that the sequence is not accessible. Species with a star symbol (*) have been sequenced in this article. TMUH: Tarbiat Modares University Herbarium, TARI: Herbarium of Research Institute of Forests and Rangelands, FUMH: Ferdowsi University of Mashhad Herbarium.

Anchusa officinalis L., AF531080 (Papini et al. 2002)/AY045703 (Bigazzi et al. 2002); Anchusa thessala Boiss. and Spruner, AF531084 (Papini et al. 2002)/AF530599 (selvi et al. 2002); Arnebia guttata Bge., EF199862 (Hu et al. 2006)/EF199872 (Hu et al. 2006); Arnebia euchroma (Royle) Johnst., EF199848 (Hu et al. 2006)/EF199874 (Hu et al. 2006); Arnebia szechenyi Kanitz, EF199863 (Hu et al. 2006)/EF199871 (Hu et al. 2006); Asperugo procumbens L., AB758290 *(Iran: Kazempour, 2007-3TMUH)/AB758320 *(Iran: Kazempour, 2007-3TMUH); Borago officinalis L., AF091151 (Ferguson 1998)/–; Brunnera orientalis (Schenk) Johnst., AY383289 (Hilger et al. 2004)/AF530601 (Selvi et al. 2002); Buglossoides purpurocaerulea (L.) Johnst., AJ555897 (Langstrom and Chase 2002)/–; Cerinthe minor L., AB758291 *(Iran: Kazempour, 2008-6 TMUH)/L43203 (Bohle et al. 1996); Cryptantha flavoculata Payson., AF091154 (Ferguson 1998)/–; Cynoglossum creticum Miller, FR715303 (Selvi et al. 2011)/–; Cynoglossum officinale L., AB758292 *(Iran: Assadi, 73526TARI)/ AB758321 *(Iran: Assadi, 73526TARI); Cynoglottis barrelieri (All.) Vural and Kit, AF531081 (Papini et al. 2002)/AY045708 (Bigazzi et al. 2002); Echiochilon fruticosum Desf., EU044843 (Thomas et al. 2007)/EU044881 (Thomas et al. 2007); Echiochilon persicum (Burm.f.) Johnst., AB758293 *(Iran: Mozaffarian, 49917TARI)/AB758322 *(Iran: Mozaffarian, 49917TARI); Echium vulgare L., AJ555896 (Langstrom and Chase 2002)/–; Eritrichium canum (Benth.) Kitam., AB758294 *(Germany: cultivated in Munich Botanical Garden)/AB758323 *(Germany: cultivated in Munich Botanical Garden); Eritrichim nanum (Vill.) Schrad., AY092901 (Winkworth et al. 2002)/–; Gastrocotyle macedonica (All.) Vural and Kit, AF531086 (Papini et al. 2002)/AY045706 (Bigazzi et al. 2002); Heliotropium bacciferum Frossk., AB758295 *(Iran: Sonboli, 01TMUH)/AB758324 *(Iran: Sonboli, 01TMUH); Heterocaryum laevigatum (Kar. et Kir.) DC., AB758296 *(Iran: Faghihna and Zangooei, 25349FUMH)/AB758325 *(Iran: Faghihna and Zangooei, 25349FUMH); Heterocaryum subsessile Vatke., AB758297 *(Iran: Faghihna and Zangooei, 28193TMUH)/AB758326 *(Iran: Faghihna and Zangooei, 28193TMUH); Heterocaryum szovitsianum (Fisch. et Mey.) DC., AB758298 *(Iran: Kazempour Osaloo, 2007-5TMUH)/AB758327 *(Iran: Kazempour Osaloo, 2007-5TMUH); Heterocaryum rigidum DC., AB758299 *(Iran: Shafaghi and Borhan, 371TARI)/AB758328 *(Iran: Shafaghi and Borhan, 371TARI); Heterocaryum macrocarpum Zak., AB758300 *(Iran: Joharchi and Zangooei, 19639TMUH)/AB758329 *(Iran: Joharchi and Zangooei, 19639TMUH); Lappula barbata (M. B.) Gurke, AB564703 (Khoshsokhan et al. 2010)/AB564713 (Khoshsokhan et al. 2010); Lappula ceratophora (M. Pop.) M. Pop., AB758301 *(Iran: Mozaffarian, 58407TARI)/AB758330 *(Iran: Mozaffarian, 58407TARI); Lappula drobovi M. Pop. ex Pavl., AB758302 *(Iran: Faghihnia and Zangooei, 26636FUMH)/AB758331 *(Iran: Faghihnia and Zangooei, 26636FUMH); Lappula microcarpa (Ledeb.) Gurke, AB758303 *(Iran: Kazempour Osaloo, 2007-6TMUH)/AB758332 *(Iran: Kazempour Osaloo, 2007-6TMUH); Lappula microcarpa (Ledeb.) Gurke, AB758304 *(Iran: Assadi, 75090TARI)/–; Lappula patula (Lehm.) Ascherson ex Gurke, AB758305 *(Iran: Assadi, 73860TARI)/AB758333 *(Iran: Assadi, 73860TARI); Lappula semiglabra (Ledeb.) Gurke, AB758306 *(Iran: Mozaffarian, 42998TARI)/AB758334 *(Iran: Mozaffarian, 42998TARI); Lappula squarrosa (Retz.) Durmort., AB758307 *(Germany: cultivated in Munich Botanical Garden)/AB758335 *(Germany: cultivated in Munich Botanical Garden); Lappula sessiliflora (Boiss.) Gurke, AB564704 (Khoshsokhan et al. 2010)/AB564714 (Khoshsokhan et al. 2010); Lappula sinaica (DC.) Ascherson ex Schweinf., AB758308 *(Iran: Kazempour Osaloo, 2007-7TMUH)/AB758336 *(Iran: Kazempour Osaloo, 2007-7TMUH); Lappula spinocarpos (Forssk.) Ascherson et Kuntze, AB758309 *(Iran: Kazempour Osaloo, 2007-4TMUH)/AB758337 *(Iran: Kazempour Osaloo, 2007-4TMUH); Lepechiniella albiflora Riedl., AB758310 *(Afghanistan: Rechinger, 31424TARI)/-; Lepechiniella inconspicua (Brand) Riedl, AB758311 *(Iran: Joharchi and Ayatollahi, 10711FUMH)/AB758338 *(Iran: Joharchi and Ayatollahi, 10711FUMH); Lepechiniella persica (Boiss.) Riedl, AB758312 *(Iran: Assadi and Maassumi, 51278TARI)/AB758339 *(Iran: Assadi and Maassumi, 51278TARI); Lepechiniella wendelboi Riedl, AB758313 *(Iran: Assadi and Maassumi, 21064TARI)/AB758340 *(Iran: Assadi and Maassumi, 21064TARI); Lepechiniella wendelboi Riedl, AB758314 *(Iran: Kazempour Osaloo, 2008-7TMUH)/–; Lindelofia longiflora Baill., AJ555895 (Langstrom and Chase 2002)/–; Myosotidium hortensia (Decne) Baill, AY092902 (Winkworth et al. 2002)/–; Myosotis abyssinica Boiss. and Reuter, AY092904 (Winkworth et al. 2002)/–; Myosotis arvensis (L.) Hill., AY092908 (Winkworth et al. 2002)/–; Myosotis asiatica Schischk, AB758315 *(Iran: Serkart and Olfat, 167TARI)/AB758341 *(Iran: Serkart and Olfat, 167TARI); Myosotis australis R. Br., AY092910 (Winkworth et al. 2002)/–; Myosotis lithospermifolia (Willd.) Hornem., AY092923 (Winkworth et al. 2002)/AB758342 *(Iran: Babakhanloo and Amin, 18179TARI); Myosotis olympica Boiss., –/AB758343 *(Iran: Assadi and Shahsavari, 65961TARI); Myosotis sylvstica Her. ex Hoffm., AY092935 (Winkworth et al. 2002)/–; Omphalodes japonica Maxim., DQ320747 (Serrano et al. 2005)/–; Omphalodes kuzinskyanae Willk., AY837595 (Serrano et al. 2004)/–; Omphalodes linifolia L., AB758316 *(Germany: cultivated in Munich Botanical Garden)/AB758344 *(Germany: cultivated in Munich Botanical Garden); Omphalodes linifolia L., AY837597 (Serrano et al. 2004)/–; Omphalodes nitida Hoffmanns and Link, AY837615 (Serrano et al. 2004)/–; Onosma hookeri Nyman., EF199847(Hu et al. 2006)/–; Onosma paniculatum Bueau and French., EF199859 (Hu et al. 2006)/EF199851 (Hu et al. 2006); Onosma waltoni Duthic, EF199846 (Hu et al. 2006)/–; Paracaryum persicum (Boiss.) Boiss., AB758317 *(Iran: Kazempour Osaloo, 2007-8TMUH)/AB758345 *(Iran: Kazempour Osaloo, 2007-8TMUH); Rochelia bungei Trautv., AB564695 (Khoshsokhan et al. 2010)/AB564705 (Khoshsokhan et al. 2010); Rochelia cardiosepala Bge., AB564601 (Khoshsokhan et al. 2010)/AB564711 (Khoshsokhan et al. 2010); Rochelia disperma (L. F.) Koch., AB564698 (Khoshsokhan et al. 2010)/AB564708 (Khoshsokhan et al. 2010); Rochelia macrocalyx Bge., AB564600 (Khoshsokhan et al. 2010)/AB564710 (Khoshsokhan et al. 2010); Rochelia mirheydari Reidl et Esfandiari, AB564696 (Khoshsokhan et al. 2010)/AB564706 (Khoshsokhan et al. 2010); Rochelia persica Bge. ex. Boiss., AB564697 (Khoshsokhan et al. 2010)/AB564707 (Khoshsokhan et al. 2010); Rochelia peduncularis Boiss., AB564699 (Khoshsokhan et al. 2010)/AB564709 (Khoshsokhan et al. 2010); Solenanthus circinatus Ledeb., AB758318 *(Iran: Khoshsokhan, 2007-9TMUH)/AB758346 *(Iran: Khoshsokhan, 2007-9TMUH); Symphytum × uplandicum, AY092903 (Winkworth et al. 2002)/–; Trichodesma aucheri DC., AB758319 *(Iran: Mozaffarian, 57195TARI)/AB758347 *(Iran: Mozaffarian, 57195TARI); Tournefortia rubicunda Salzm.ex DC., EF688852 (Luebert et al. 2008)/EF688799 (Luebert et al. 2008); Trigonotis peduncularis (Trevir.) Benth., DQ320740 (Serrano et al. 2005)/–.

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Khoshsokhan Mozaffar, M., Kazempour Osaloo, S., Oskoueiyan, R. et al. Tribe Eritrichieae (Boraginaceae s.str.) in West Asia: a molecular phylogenetic perspective. Plant Syst Evol 299, 197–208 (2013). https://doi.org/10.1007/s00606-012-0715-4

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