Abstract
Previous molecular phylogenetic investigations of Apiaceae tribe Pimpinelleae have focused primarily on its largest genus Pimpinella and its closest allies. The monophyly and phylogenetic placements of five Iranian genera of the tribe have not been addressed sufficiently (Aphanopleura, Demavendia, Haussknechtia, Psammogeton, and Zeravschania). To examine relationships, a nrDNA ITS matrix including 169 accessions representing 49 genera of Apiaceae (including 10 Iranian taxa not analyzed previously) and a cpDNA rps16 matrix containing 37 accessions representing 24 genera of the family, representing the greatest sampling to date of the aforementioned genera, were subjected to phylogenetic analyses using Bayesian inference and maximum parsimony methods. The trees obtained showed a close affinity among the examined species of Aphanopleura, Psammogeton and several species of Trachyspermum. Neither Aphanopleura nor Psammogeton resolved as monophyletic, and A. leptoclada allied with Pimpinella. The genera Demavendia, Haussknechtia and Zeravschania also comprised a well-supported clade, with Demavendia and Haussknechtia (in the ITS trees) arising from within a paraphyletic Zeravschania. To recognize monophyletic genera, one new combination is proposed in Pimpinella and six new combinations are proposed in Psammogeton. A broader circumscription of Zeravschania to include Demavendia and Haussknechtia may also be warranted, but must await further study.
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Acknowledgements
S.S.M. is grateful to the University of Tehran and the Research Institute of Forests and Rangelands in Tehran for supporting her work. We are grateful to the directors and curators of the herbaria TARI and TUH for their generous assistance in providing plant materials and photographs for this study. S.Z. also appreciates the financial support provided by Alexander Humboldt Stiftung (Germany) and the University of Tehran. Lastly, we thank three anonymous reviewers and Deborah S. Katz-Downie for their valuable constructive comments on early drafts of the manuscript.
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Appendices
Appendix
Accessions of Pimpinelleae and other taxa of Apiaceae subfamilies Apioideae and Saniculoideae included in the study. Previously published nrDNA ITS and rps16 sequences are represented herein by GenBank accession numbers only. The first GenBank number following a species name refers to ITS and the second for rps16, except for those instances where there are three numbers and these refer to separate ITS1, ITS2, and rps16 sequences. A single GenBank number often refers to ITS but in a few instances it refers to rps16. The number “1” or “2” added to the species name coincides with their placement in Figs. 2 and 3. For those species with newly generated ITS and rps16 sequences, voucher information including species name, geographical origin, collector(s), vouchers (herbarium: TARI, Tehran Ariamehr Research Institute) is provided. Species names in brackets follow the taxonomic treatment proposed in the present study.
Aciphylla glacialis, KF160671; Aegopodium tribracteolatum, AY581783; Aethusa cynapium, AF110539; Aletes macdougalii, KF619604; Ammi trifoliatum, KJ473904; Ammodaucus leucotrichus, KT347642, KT347831; Annesorhiza macrocarpa, DQ368835; Anthriscus sylvestris, KT347715, KT347879; Aphanopleura capillifolia [Psammogeton capillifolium], DQ516368; Aphanopleura breviseta [Psammogeton brevisetum], Iran, Prov. Azarbaijan, ca 2 km from Tabriz to Bababaghi,1400 m a. s. l., Mozaffarian 71,290 (TARI), MZ687311, MZ706974; Aphanopleura leptoclada [Pimpinella leptoclada], Iran, Prov. Yazd, Mehriz to Tezerjan, Sand dunes around the roads, 1500 m a. s. l., Mozaffarian 77,439 (TARI), MZ687308, MZ706975; Aphanopleura trachysperma1, AF008629; Aphanopleura trachysperma2, Iran, Prov. West Azarbaijan, Maku, Shut, SE of Sufi village, 1070 m a. s. l., Mozaffarian 94,396 (TARI), MZ687310, MZ706976; Apium graveolens, AH003471; Arafoe aromatica1, AF077874; Arafoe aromatica2, U78383, U78443; Astrodaucus orientalis, MG098986, AF123748; Bunium allioides, JX312805; Bunium setaceum, EF544477; Bupleurum microcephalum, GU269882; Carum carvi, KF454471, FJ385182; Chamaesium paradoxum, EU236161; Choritaenia capensis, DQ368842; Conium maculatum, AF110546; Coriandrum sativum, HQ377205; Cryptotaenia africana1, DQ516371; Cryptotaenia africana2, DQ516370; Cymbocarpum wiedemannii, GU291352; Demavendia pastinacifolia, AY911857, AY911863; Demavendia pastinacifolia, Iran, Prov. Tehran, W Tehran, Suleghun valley, 1500–2000 m a. s. l., Assadi & Mozaffarian 32,621 (TARI), MZ687312, MZ706977; Ducrosia flabellifolia, DQ427051; Echinophora chrysantha, AF077883; Erigenia bulbosa, MG218514; Eryngium proteiflorum, EU070531; Ferula grigoriewii, KJ660784, KJ698392; Frommia ceratophylloides, DQ647630; Galagania neglecta, HM229391; Haussknechtia elymaitica1, EU169273; Haussknechtia elymaitica2, AY911859, AY911865; Heteromorpha involucrata, DQ368853; Komarovia anisosperma, AF077897; Lecokia cretica, EU169294; Lichtensteinia trifida, EU434683; Muretia lutea, DQ516359; Nothosmyrnium japonicum1, DQ516367, FJ385211; Nothosmyrnium japonicum2, EU236179; Oedibasis tamerlanii, HM229403; Opopanax armeniacus, Iran, Prov. East Azarbayejan: Mianeh, Bozgoush Mount. Neshagh village, 1880 m a. s. l., 2007, Mozaffarian 93,480 (TARI), MZ687309; Orlaya kochii, AH003483, AF123733; Ormopterum turcomanicum, GQ379313; Petroselinum crispum, GQ148800, AF110544; Phellolophium madagascariense1, DQ647627, KJ173509; Phellolophium madagascariense2, KJ173483, KJ173508; Physospermopsis cuneata, FJ385055, FJ385221; Physospermopsis shaniana, EU236192, FJ385225; Pimpinella acuminata, EU236193, FJ385226; Pimpinella affinis, AY581780; Pimpinella alismatifolia, FM986448; Pimpinella anisum1, KR150177, KJ173510; Pimpinella anisum2, KR150177; Pimpinella armena, KX982510; Pimpinella arguta, JF831512; Pimpinella aromatica, AY581784; Pimpinella aurea, AY581785; Pimpinella austriaca, KX982512; Pimpinella betsileensis1, DQ647626, KJ173511; Pimpinella betsileensis2, KJ173488; Pimpinella brachycarpa, AY548230; Pimpinella brachystyla, GQ379270; Pimpinella buchananii, FM986455; Pimpinella caffra, FM986447; Pimpinella candolleana1, MH117649, FJ385227; Pimpinella candolleana2, MH117649; Pimpinella cappadocica, AY581786; Pimpinella caudata, JF831514; Pimpinella chungdienensis, JF831515; Pimpinella confuse, KX982508; Pimpinella coriacea, JF831516; Pimpinella corymbosa, AY581787; Pimpinella cretica, AY581788; Pimpinella diversifolia1, JF831517; Pimpinella diversifolia2, KF806585; Pimpinella eriocarpa, AY581790; Pimpinella espanensis, MH377861; Pimpinella fargesii, JF831518; Pimpinella favifolia1, FM986458; Pimpinella favifolia2, FM986453; Pimpinella flabellifolia, AY581791; Pimpinella flaccida, JF831519; Pimpinella henryi, EU236195; Pimpinella heyneana, GQ379276; Pimpinella huillensis1, FM986454; Pimpinella huillensis2, FM986443; Pimpinella insignis, AY941280, AY941308; Pimpinella isaurica, AY581792; Pimpinella khayyamii, KX982523; Pimpinella kingdon-wardii, JF831520; Pimpinella kotschyana, DQ516373; Pimpinella krookii, FM986445; Pimpinella kyimbilaensis, FM986452; Pimpinella ledermannii, FM986457; Pimpinella lutea, DQ516374; Pimpinella major1, MH377862; Pimpinella major2, KX982513; Pimpinella nigra, KX982509; Pimpinella niitakayamensis, DQ516375; Pimpinella nudicaulis, AY581794; Pimpinella olivieri1, KX982506; Pimpinella olivieri2, KX922690; Pimpinella oliverioides, AY581795; Pimpinella oreophila, FM986450; Pimpinella peregrina1, AY581797; Pimpinella peregrina2, AY581797; Pimpinella peucedanifolia1, AY581798; Pimpinella peucedanifolia2, KX982524; Pimpinella procumbens, MH377863; Pimpinella puberula, AY581799; Pimpinella purpurea1, JF831521; Pimpinella purpurea2, EU236197; Pimpinella rhodantha1, KX982511; Pimpinella rhodantha2, KX982511; Pimpinella rhomboidea, JF831522; Pimpinella rigidistyla, FM986459; Pimpinella rockii1, JF831523; Pimpinella rockii2, FJ385057; Pimpinella rubescens, JF831524; Pimpinella saxifraga1, MH377864, DQ133876; Pimpinella saxifraga2, AY581801; Pimpinella serbica, KX982507; Pimpinella siifolia1, MH377868; Pimpinella siifolia2, MH377867; Pimpinella sintenisii, AY581802; Pimpinella smithii1, GQ379272, FJ385230; Pimpinella smithii2, JF831526; Pimpinella thellungiana, JF831527; Pimpinella tibetanica, JF831528; Pimpinella tragium1, MH377869; Pimpinella tragium2, MH377869; Pimpinella transvaalensis, FM986449; Pimpinella trifurcata, FM986446; Pimpinella valleculosa, JF831529; Pimpinella villosa, MH377871; Pimpinella yunnanensis1, JF831530, FJ385231; Pimpinella yunnanensis2, JF831530; Pleurospermum rivulorum, HQ824798; Postiella capillifolia, DQ422829, DQ422848; Prangos eriantha, MT250532; Prangos ferulacea, KX982519; Prangos lophoptera, KU168369; Psammogeton biternatum, AF164839, AF164864; Psammogeton canescens, MG827069; Psammogeton lamondiae, MG827073; Psammogeton ranunculifolius, MG920277; Psammogeton sp., MG827076; Psammogeton stocksii, Iran, Prov. Baluhestan, Between Iranshahr and Bam, Bazman, 1200 m a. s. l., Assadi 23,009 (TARI), MZ687313, MZ706978; Pyramidoptera cabulica, AF008631; Sanicula epipactis, EU169013, EU168959; Selinum broteri, AY179029; Smyrnium olusatrum, MK050086; Trachydium simplicifolium, FJ385067; Trachyspermum ammi, Iran, Prov. Baluchistan: Iranshahr to Chahbahar, Pole Jaligur over the Sarbay river, 300 m a. s. l., Mozaffarian 74,383 (TARI), MZ687314, MZ706979; Trachyspermum anethifolium [Psammogeton anethifolium], MG827079; Trachyspermum copticum, Iran, Prov. Baluchistan: Khash to Iranshahr, Dashte Abkhan, 1400 m a. s. l., Mozaffarian 72,711 (TARI), MZ687315, MZ706980; Trachyspermum microcarpum [Psammogeton microcarpum], MG920276; Trachyspermum paktianum [Psammogeton paktianum], MG827078; Trachyspermum papillare [Psammogeton papillare], MG920275; Trachyspermum roxburghianum [Psammogeton involucratum], MG827080; Trachyspermum scaberulum1, JF831531, FJ385258; Trachyspermum scaberulum2, EU236215; Trachyspermum triradiatum, EU236216, FJ385259; Zeravschania aucheri, AY911860, AY911866; Zeravschania membranacea1, AY911862, AY911868; Zeravschania membranacea2, Iran, Prov. Khorassan: Mashhad, Karde. All village, 1460 m a. s. l., Mozaffarian 87,007 (TARI), MZ687317, MZ706982; Zeravschania pauciradiata, Iran, Prov. East Azarbaijan, Siahrud to Kakaleh, near to Kalaleh, 390 m a. s. l., Ajani 1584 (TARI), MZ687316, MZ706981; Zeravschania regeliana, AY911861, AY911867.
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Online Resource 1. The alignment of nrDNA ITS sequences which has been used to reconstruct the ITS trees obtained in this study.
Online Resource 2. The alignment of plastid rps16 intron sequences which has been used to reconstruct the rps16 trees obtained in this study.
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Mousavi, S.S., Mozaffarian, V., Mummenhoff, K. et al. Systematics of the Iranian genera Aphanopleura, Demavendia, Haussknechtia, Psammogeton, and Zeravschania (Apiaceae tribe Pimpinelleae). Plant Syst Evol 308, 2 (2022). https://doi.org/10.1007/s00606-021-01792-x
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DOI: https://doi.org/10.1007/s00606-021-01792-x