Abstract
The article presents a find of Carex ×favratii (Carex echinata × C. paniculata) in Romania, whose genetic relationships with its putative parents was tested using AFLP and microsatellite data. The putative hybrid was genetically intermediate between the parental taxa, i.e. C. echinata and C. paniculata. A morphological analysis shows ten intermediary characters, six ones closer to C. echinata, five closer to C. paniculata, and three unique, whereas the perigynium and achene characters could not be evaluated due to hybrid sterility (with exception of the type of perigynium). The morphological features supported by molecular markers strongly support the hybrid nature of the collected plants. The results provide evidence that a hybrid between these species really occurs in nature.
References
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Korpelainen, H., Virtanen, V., Kostamo, K. & Väre, H. (2010) Hybridization and introgression in Carex aquatilis and C. paleacea. Plant Systematics and Evolution 287: 141–151. https://doi.org/10.1007/s00606-010-0307-0
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?epka, R., Veselá, P. & Mrá?ek, J. (2014) Are there hybrids between Carex flacca and C. tomentosa in the Czech Republic and Slovakia? Preslia 86: 367–379.
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Smith, T.W. & Waterway, M.J. (2008) Evaluating species limits and hybridization in the Carex complanata complex using morphology, amplified fragment length polymorphisms, and restriction fragment analysis. Botany 86: 809–826. https://doi.org/10.1139/B08-031
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Vos, P., Hogers, R., Bleeker, M., Reijans, M., van de Lee, T., Hornes, M., Frijters, A., Pot, J., Peleman, J., Kuiper, M. & Zabeau, M. (1995) AFLP: a new technique for DNA fingerprinting. Nucleic Acids Research 23: 4407–4414. https://doi.org/10.1093/nar/23.21.4407
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Wi?c?aw, H. & Koopman, J. (2013) Numerical analysis of morphology of natural hybrids between Carex hostiana and the members of Carex flava agg. (Cyperaceae). Nordic Journal of Botany 31: 464–472. https://doi.org/10.1111/j.1756-1051.2013.00095.x
Wi?c?aw, H. & Wilhelm, M. (2014) Natural Hybridization within the Carex flava Complex (Cyperaceae) in Poland: Morphometric Studies. Annales Botanici Fennici 51: 129–147. https://doi.org/10.5735/085.053.0101
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Blackstock, N. & Ashton, P.A. (2010) Genetic markers and morphometric analysis reveal past hybridization and introgression in putative Carex flava L. s. str. (Cyperaceae) hybrid populations. Plant Systematics and Evolution 287: 37–47. https://doi.org/10.1007/s00606-010-0287-0
Boott, F.M.B. (1838) Carex recta Boott in Hooker W.J. Flora Boreali-Americana. II. H.G. Bohn, Covent Garden, London 328 pp.
Bugg, C., Smith, C., Blackstock, N., Simpson, D. & Ashton, P.A. (2013) Consistent and variable leaf anatomical characters in Carex (Cyperaceae). Botanical Journal of the Linnean Society 172: 371–384. https://doi.org/10.1111/boj.12029
Catling, P.M. (1993) Carex castanea × Carex debilis, a new natural hybrid from Ontario. Rhodora 95: 129–136.
Cayouette, J. & Catling, P.M. (1992) Hybridization in the genus Carex with special reference to North America. Botanical Review 58: 351–440. https://doi.org/10.1007/BF02858773
Christ, H. (1885) Nouveau catalogue des Carex d´Europe. Bulletin de la Société Royale de Botanique de Belgique 24: 10–20.
Christ, H. (1889) Second appendice au nouveau catalogue des Carex d’Europe. Bulletin de la Société Royale de Botanique de Belgique 28: 165–167.
Dumortier, B.C.J. (1827) Florula Belgica. Casterman, Tournay, 172 pp.
Escudero, M., Eaton, D.A.R., Hahn, M. & Hipp, A.L. (2014) Genotyping-by-sequencing as a tool to infer phylogeny and ancestral hybridization: A case study in Carex (Cyperaceae). Molecular Phylogenetics and Evolution 79: 359–367. https://doi.org/10.1016/j.ympev.2014.06.026
Escudero, M., Hipp, A.L. & Luceño, M. (2010) Karyotype stability and predictors of chromosome number variation in sedges: A study in Carex section Spirostachyae (Cyperaceae). Molecular Phylogenetics and Evolution 57: 353–363. https://doi.org/10.1016/j.ympev.2010.07.009
Ford, B.A., Ghazvini, H., Naczi, R.F.C. & Starr, J.R. (2012) Phylogeny of Carex subg. Vignea (Cyperaceae) based on Amplified Fragment Length Polymorphism and nrDNA Data. Systematic Botany 37: 913–925. https://doi.org/10.1600/036364412X656464
Ford, B.A., Ball, P.W. & Ritland, K. (1993) Genetic and macromorphologic evidence bearing on the evolution of members of Carex section Vesicariae (Cyperaceae) and their natural hybrids. Canadian Journal of Botany 71: 486–500. https://doi.org/10.1139/b93-053
Fries, E.M. (1945) Summa vegetabilium Scandinaviae 1. Uppsaliae. 258 pp.
Gray, A. (1848) Manual of Botany of the Northern United States. J. Munroe, Boston & J.Chapman, London. 710 pp. https://doi.org/10.5962/bhl.title.128844
Heathcote, C.A., Davies, M.S. & Etherington, J.R. (1987) Phenotypic flexibility of Carex flacca Schreb. Tolerance of soil flooding by populations from contrasting habitats. New Phytologist 105: 381–391. https://doi.org/10.1111/j.1469-8137.1987.tb00875.x
Hendrichs, M., Michalski, S., Begerow, D., Oberwinkler, F. & Hellwig, F.H. (2004) Phylogenetic relationship in Carex, subgenus Vignea (Cyperaceae), based on ITS sequences. Plant Systematics and Evolution 246: 109–125. https://doi.org/10.1007/s00606-004-0127-1
Hipp, A.L., Rothrock, P.E., Reznicek, A.A. & Berry, P.E. (2007) Chromosome Number Changes Associated with Speciation in Sedges: a Phylogenetic Study in Carex section Ovales (Cyperaceae) Using AFLP Data. Aliso: A Journal of Systematic and Floristic Botany 23: 193–203. https://doi.org/10.5642/aliso.20072301.14
Hipp, A.L., Rothrock, P.E. & Roalson, E.H. (2009) The evolution of chromosome arrangements in Carex (Cyperaceae). Botanical Review 75: 96–109. https://doi.org/10.1007/s12229-008-9022-8
Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (1998) Online updates version of Index Herbariorum Part I: The herbaria of the World, 8<sup>th</sup> edn. [http://www.nybg.org/bsci/ih/ih.html]
Jiménez-Mejías, P., Escudero, M., Guerra-Cárdenas, S., Lye, K.A. & Luceño, M. (2011) Taxonomic delimitation and drivers of speciation in the Ibero-North African Carex sect. Phacocystis river-shore group (Cyperaceae). American Journal of Botany 98: 1855–1867. https://doi.org/10.3732/ajb.1100120
Jiménez-Mejías, P., Luceño, M., Wilson, K.L., Waterway, M.J. & Roalson, E.R. (2016) Clarification of the use of the terms perigynium and utricle in Carex L. (Cyperaceae). Systematic Botany 41: 519–528. https://doi.org/10.1600/036364416X692488
Kaplan, Z., Danihelka, J., Št?pánková, J., Ekrt, L., Chrtek, J. jr., Zázvorka, J., Grulich, V., ?epka, R., Pran?l, J., Duchá?ek, M., Kúr, P., Šumberová, K. & Br?na, J. (2016) Distributions of vascular plants in the Czech Republic. Part 2. Preslia 88: 229–322. https://doi.org/10.23855/preslia.2017.115
Kaplan, Z., Danihelka, J., Koutecký, P., Šumberová, K., Ekrt, L., Grulich, V., ?epka, R., Hroudová, Z., Št?pánková, J., Dvo?ák, V., Dan?ák, M., D?evojan, P. & Wild, J. (2017) Distributions of vascular plants in the Czech Republic. Part 4. Preslia 89: 115–201. https://doi.org/10.23855/preslia.2017.115
King, M.G. & Roalson, E.H. (2009) Isolation and characterisation of 11 microsatellite loci from Carex macrocephala (Cyperaceae). Conservation Genetics 10: 531–533. https://doi.org/10.1007/s10592-008-9558-5
Koopman, J. (2011) Carex Europaea. The genus Carex L. (Cyperaceae) in Europe, 1. Margraf Publishers, Weikersheim. 726 pp.
Korpelainen, H., Virtanen, V., Kostamo, K. & Väre, H. (2010) Hybridization and introgression in Carex aquatilis and C. paleacea. Plant Systematics and Evolution 287: 141–151. https://doi.org/10.1007/s00606-010-0307-0
Košnar, J., Štech, M. & Koutecký, P. (2012) Environmental control of clonal growth in Carex nigra: What can be masked under the name Carex nigra subsp. juncella in the Czech Republic? Flora 207: 294–302. https://doi.org/10.1016/j.flora.2011.12.001
Kunth, C.S. (1837) Enumeratio Plantarum (Cyperographia synoptica) 2. Stutgardiae, Tubingae. 591 pp.
Linnaeus, C. (1753) Species Plantarum. Impensis Laurentii Salvii. Holmiae. 560 pp.
Linnaeus, C. (1755) Centuria I Plantarum. L. M. Ho?jer, Regio Acad. Typogr. Uppsaliae. 35 pp.
Linnaeus, C. (1767): Mantissa Plantarum. Generum editionis VI. et specierum editionis II. Impensis Direct. Laurentii Salvii. Holmiæ. 142 pp. https://doi.org/10.5962/bhl.title.119809
Meinshausen, K.F. (1901) Die Cyperaceen der Flora Russlands. Trudy botani?eskogo sada 18: 221–415.
Murray, J.A. (1773) Prodromus Designationis Stirpium Gottingensium cum Figuris Aeneis. Gottingae. 252 pp.
Nakamatte, E. & Lye, K.A. (2007) AFLP-based differentiation in north Atlantic species of Carex sect. Phacocystis. Nordic Journal of Botany 25: 318–328. https://doi.org/10.1111/j.0107-055X.2007.00116.x
Ohsako, T. & Yamane, K. (2007) Isolation and characterization of polymorphic microsatellite loci in Asiatic sand sedge, Carex kobomugi Ohwi (Cyperaceae). Molecular Ecology Notes 7: 1023–1025.
Peakall, R. & Smouse, P.E. (2012) GenAlEx 6.5: genetic analysis in Excel. Population genetic software for teaching and research—an update. Bioinformatics 28: 2537–2539. https://doi.org/10.1093/bioinformatics/bts460
Rieseberg, L.H. (1995) The role of hybridization in evolution: old wine in new skins. American Journal of Botany 82: 944–953. https://doi.org/10.1002/j.1537-2197.1995.tb15711.x
Roalson, E.H., Jiménez-Mejías, P., Hipp, A.L., Benítez-Benítez, C., Bruederle, L.P., Chung, K.-S., Escudero, M., Ford, B.A., Ford, K., Gebauer, S., Gehrke, B., Hahn, M., Hayat, M.Q., Hoffmann, M.H., Jin, X.-F., Kim, S., Larridon, I., Léveillé-Bourret, É., Lu, Y.-F., Luceño, M., Maguilla, E., Márquez-Corro, J.I., Martín-Bravo, A., Masaki, T., Míguez, M., Naczi, R.F.C., Reznicek, A.A., Spalink, D., Starr, J.R., , Villaverde, T., Waterway, M.J., Wilson, K.L. & Zhang, S.-R. (2021) A framework infrageneric classification of Carex (Cyperaceae) and its organizing principle. Journal of Systematics and Evolution 59: 726–762. https://doi.org/10.1111/jse.12722
?epka, R., Veselá, P. & Mrá?ek, J. (2014) Are there hybrids between Carex flacca and C. tomentosa in the Czech Republic and Slovakia? Preslia 86: 367–379.
Schreber, J.C.D. (1771) Spicilegium florae lipsicae. Dykiano, Lipsiae. 148 pp. https://doi.org/10.5962/bhl.title.6360
Smith, T.W. & Waterway, M.J. (2008) Evaluating species limits and hybridization in the Carex complanata complex using morphology, amplified fragment length polymorphisms, and restriction fragment analysis. Botany 86: 809–826. https://doi.org/10.1139/B08-031
Stenström, A., Jonsson, B.O., Jónsdóttir, I.S., Fagerström, T. & Augner, M. (2001) Genetic variation and clonal diversity in four clonal sedges (Carex) along the Arctic coast of Eurasia. Molecular Ecology 10: 497–513. https://doi.org/10.1046/j.1365-294x.2001.01238.x
Stenström, A., Jónsdóttir, I.S. & Augner, M. (2002) Genetic and environmental effects on morphology in clonal sedges in the Eurasian Arctic. American Journal of Botany 89: 1410–1421. https://doi.org/10.3732/ajb.89.9.1410
Urbanek, L. (1998) Morphometric differentiation of Carex ligerica Gay in Poland. Acta Societatis Botanicorum Poloniae 67: 263–268. https://doi.org/10.5586/asbp.1998.034
Vegetti, A.C. (2002) Typological reinterpretation of the inflorescences in Cariceae (Cyperaceae). Phyton 42: 159–168.
Villars, D. (1787) Histoire des Plantes de Dauphiné. Vol. 2. Société royale de médecine, Grenoble. 690 pp. https://doi.org/10.5962/bhl.title.116232
Volkova, P.A., Shipunov, A.B., Elven, R. & Brochmann, C. (2008) The seashore sedges of the Russian Kola Peninsula: how many species? Flora 203: 523–533. https://doi.org/10.1016/j.flora.2007.09.004
Vos, P., Hogers, R., Bleeker, M., Reijans, M., van de Lee, T., Hornes, M., Frijters, A., Pot, J., Peleman, J., Kuiper, M. & Zabeau, M. (1995) AFLP: a new technique for DNA fingerprinting. Nucleic Acids Research 23: 4407–4414. https://doi.org/10.1093/nar/23.21.4407
Wahlenberg, G. (1803): Inlending til Caricographien. Kongl. Veteskamps Academiens Nya Handlingar 24: 225–329.
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