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Phylogeny, karyotype evolution and taxonomy of Crocus series Verni (Iridaceae)

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Abstract

The taxonomically complicated Crocus series Verni is characterized by high intra- and interspecific variability of karyotypes (2n = 8–23). With the aim to get more insights into complex karyotype evolution and to clarify the taxonomy of this group, we combined morphological (twelve characters), molecular (chloroplast DNA: trnL–trnF, ndhF; nuclear DNA: ITS, pCOSAt103) and karyological analyses. Samples of different populations of C. etruscus, C. ilvensis, C. kosaninii, C. tommasinianus, C. vernus sensu lato and C. longiflorus (series Longiflori) were analyzed. Quantitative karyotype parameters were calculated for all taxa involved based on the available literature. For the taxon traditionally known as C. vernus, the analyses suggest that it should be split in five species: C. heuffelianus, C. neapolitanus, C. neglectus sp. nov., C. siculus and C. vernus. The comparison of genome total haploid lengths suggests that in the evolution of the group polyploidization only played a role within the C. vernus species complex, where we also detected two hybridization events. In all other taxa, chromosome evolution is probably characterized by chromosome fusions and fissions, sometimes affecting the entire haploid chromosome set. Comparative cytogenetics of the group indicates that series Verni is subject to a peculiar type of unequal change in chromosome size, i.e., that not both chromosome arms gain or lose equally in DNA content. As a taxonomic consequence of our study, series Verni is newly circumscribed, now including the autumn-flowering C. longiflorus and excluding C. baytopiorum.

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Acknowledgments

We thank B. Kränzlin and P. Oswald for expert technical help in the lab, F. Bartolucci for logistic help in collecting material of C. neapolitanus in Abruzzo, G. Gestri and B. Pierini for collecting material of C. neglectus, L. Bernardo and D. Gargano for providing Crocus materials conserved in CLU, Thomas Huber and Dirk Schnabel for providing Crocus material of their living collection and for measuring morphological parameters for these specimen, H. Kerndorff for his comments on the manuscript, Z. Barina for providing images of Crocus heuffelianus in BP, G. Domina for useful advices about Tineo’s life and calligraphy. The Ministry of Education, Science and Technological Development of the Republic of Serbia supported this research through Grant 173030 “Plant biodiversity of Serbia and the Balkans–assessment, sustainable use and protection” and the German Research Foundation (DFG) through Grant BL462/7.

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Authors and Affiliations

  1. Leibniz Institute of Plant Genetics and Crop Research (IPK), 06466, Gatersleben, Germany

    Dörte Harpke & Frank R. Blattner

  2. Unit of Botany, Department of Biology, University of Pisa, via Luca Ghini 13, 56126, Pisa, Italy

    Angelino Carta & Lorenzo Peruzzi

  3. Faculty of Biology, Institute of Botany and Botanical Garden, University of Belgrade, Takovska 43, 11000, Belgrade, Serbia

    Gordana Tomović

  4. Department of Biology and Ecology, Faculty of Natural Sciences, University of Niš, Višegradska 33, 18000, Niš, Serbia

    Vladimir Ranđelović & Novica Ranđelović

  5. German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, 04103, Leipzig, Germany

    Frank R. Blattner

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  1. Dörte Harpke
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Correspondence to Dörte Harpke.

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606_2014_1074_MOESM1_ESM.tif

Phylogenetic tree of series Verni derived from a Bayesian analysis of combined sequences of the ITS, trnL–trnF and ndhF regions (24 sequences, 2026 characters). Posterior probabilities are given at the nodes (TIFF 12634 kb)

Appendix. Sources of the studied material with abbreviation for populations used in text and figures in the following order: Taxon: Acronym: Origin, herbar voucher. Order of GenBank accession numbers for voucher of outgroups: trnL–trnF, ndhF, ITS (ITS1 + 5.8SrDNA + ITS2), pCOSAt103;—denotes lacking sequence

Appendix. Sources of the studied material with abbreviation for populations used in text and figures in the following order: Taxon: Acronym: Origin, herbar voucher. Order of GenBank accession numbers for voucher of outgroups: trnL–trnF, ndhF, ITS (ITS1 + 5.8SrDNA + ITS2), pCOSAt103;—denotes lacking sequence

C. neglectus 2n = 16, PE: Italy, Monte Pelato (Livorno), Tuscany, PI s.n., PI: Italy, Monte Pisano (Lucca), PI s.n.; C. neapolitanus 2n = 8, A: Italy, Altopiano delle Rocche (L’Aquila), Abruzzo, PI s.n.; AII: Italy, Collarmele (L’Aquila), Abruzzo, PI s.n.; C. siculus 2n = 8, Sic: Italy, Nebrodi (Messina), Sicily, CLU 19778, 19779; C. vernus, 2n = 8, VER: Switzerland, Ticino, Airolo, San Gottardo area, 1100-1500 m a.s.l. (locus classicus), PI s.n.; C. ilvensis 2n = 8, Fi: Italy, Capanne, Filicaia (Elba Island), Tuscany, PI s.n.; C: Italy, Capanne, Colle Grottaggia (Elba Island), Tuscany, PI, s.n.; M: Italy, Capanne, Monte Maolo (Elba Island), Tuscany, PI s.n.; N: Italy, Capanne, Valle Nivera (Elba Island), Tuscany (locus classicus), PI s.n.; C. etruscus 2n = 8, S: Italy, Sassetta, Tuscany, PI s.n.; SS: Italy, Sassetta 2nd locality, Tuscany, PI s.n.; CA: Italy, Campiglia Marittima, Tuscany, PI s.n.; F: Italy, Salita del Filetto, Tuscany (locus classicus), PI s.n.; P: Italy, Punta Ala, Tuscany, PI s.n.; C. tommasinianus 2n = 16, E: Serbia, Krusevac, village Lazarevac, C Serbia BEOU 31678, E: Serbia, Soko Banja, village Resnik, E Serbia BEOU 31679; C. kosaninii, 2n = 14, R: Serbia, Mt Rujan, Planina, S Serbia, BEOU 31680, S: Serbia, Leskovac, village Strojkovce, SE Serbia, BEOU 31681; C. heuffelianus, 2n = 22?, W Serbia: Mt Zlatibor, Tornik peak, 1450 m a.s.l., BEOU 31682; C. longiflorus, 2n = 28, T: Italy, Tortora (Cosenza), 350 m, CLU 12508, G: Italy, Grotta della Manca (Cosenza), 1250 m a.s.l., CLU 13099, Ne: Italy, Nebrodi Range, Sicilia, 1050 m a.s.l., GAT 23048, EM: Italy, Luciano 680-1000 m a.s.l., EM1057, GAT 23049.

Outgroups: C. asumaniae B.Mathew and T.Baytop, cult., GAT 7207, -, -, HE801126,-; C. banaticus J.Gay, cult., GAT 7285, -, -, HE801147,-; C. baytopiorum B.Mathew, HKEP1014, Turkey, GAT 7187, -, -, HE663994,-; C. cambessedesii J.Gay, Spain, GAT 7211, -, -, a: HE801098 d: HE801104,.-; C. cartwrightianus Herb., Greece, B/DB 4304, -, -, HG518198; C. cartwrigthianus var. creticus Herb., Greece, MAIC 1991, -, -, HE801160, -; C. corsicus Vanucchi ex Maw, France, Corsica, GAT 7252; -, -,a: HE801091 b: HE801097, -; C. goulimyi Turrill, cult., GAT 7217, -, -, HE801130, b: HE663874 efgc: HE663876; C. hadriaticus Herb, Greece, ATH 1532, -, -, HE864270,-; C. imperati Ten., Italy, GAT 7221, -, -, HE801131,-; C. ligusticus Mariotti, Italy, GAT 7226, -, -, HE801167,-; C. malyi Vis., Croatia, HK293, GAT 7259, -, -, HE801170; C. mathewii Kernd. and Pasche, Turkey, GAT 7189, -, -, HE801089,-; C. moabiticus Bornm. and Dinsm. ex Bornm., Jordan, HKEP9942, -, -, HG518196,-; C. nudiflorus Sm. Spain, HKEP1231, -, -, HG518194,-; C. oreocreticus B.L.Burtt, Greece, MAIC 3674, -, -, HE801161; C. pallasii Goldb. subsp. pallasii, cult., GAT 7230, -, -, HG518197,-; C. pallasii Turkey, HKEP1033A, GAT 7175, -, -, HE664002,-; C. pelistericus Pulević, Greece, B 10 0355313, -, -, HE664020,-; C. sativus L., cult., GAT 7277, -, -, HE801144,-; C. scardicus Košanin, Macedonia, B 10 0112048, -, -, HE663961,-; C. scharojanii Rupr., Russia, Caucasus, 2570 m a.s.l., GAT 23083, HG518229, HG518260, -,-; C. serotinus Salisb. Spain, HKEP1124, -, -, HG518195,-; C. serotnius subsp. salzmannii B.Mathew, Morocco, GAT 7276, -, -, HE801125, a: HE663913 b: HE663914; C. thomasii Ten., Italy, CLU 13102, -, -, HE801118,-; Romulea bulbocodium (L.) Sebast. and Mauri, Turkey, HKEP1108, GAT 7481, -, -, HE664008,-; Romulea ramiflora Ten., Turkey, HKEP1115, GAT 7480, -, -, HE664012,-.

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Harpke, D., Carta, A., Tomović, G. et al. Phylogeny, karyotype evolution and taxonomy of Crocus series Verni (Iridaceae). Plant Syst Evol 301, 309–325 (2015). https://doi.org/10.1007/s00606-014-1074-0

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