Abstract
Heliosperma includes about 20 taxa which are divided into alpestre, macranthum and pusillum clades based on nuclear and plastid data. Based on microstructural qualitative and quantitative seed characters of H. macranthum, H. alpestre and of nine Balkan species, we found that the species assigned to the pusillum clade are highly variable and differ from H. alpestre and H. pusillum subsp. pusillum. Two low elevation species (H. retzdorffianum and H. nikolicii) were clearly separated from the H. pusillum group, which includes alpine taxa. Newly described characters of the seeds, e.g. two hilum chambers, two rows in crest and black colour could be treated as characteristic for lowland endemics. Microstructural differences in seed character of Heliosperma species seems to be mainly driven by ecological factors resulting from the elevation gradients. Pollen is undifferentiated between species, its viability (stainability) is high, chromosome number is 2n = 26, which differs from the number reported so far (2n = 24).
References
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Cobon, A.M. & Murray, B.G. (1983) Unstable B chromosomes in Silene maritima With. (Caryophyllaceae). Botanical Journal of the Linnean Society 87: 273–283. https://doi.org/10.1111/j.1095-8339.1983.tb00995.x
Degen, A. (1937) Flora velebitica, Vol. 2. Verlag der Ungar. Akademie der Wissenschaften, Budapest, 660 pp.
EC (European Commission) (2013) Interpretation Manual of European Union Habitats. EUR 28: 1–144.
Frajman, B. & Rabeler, R.K. (2006) Proposal to conserve the name Heliosperma against Ixoca (Caryophyllaceae, Sileneae). Taxon 55 (3): 806–807. https://doi.org/10.2307/25065663
Frajman, B. & Oxelman, B. (2007) Reticulate phylogenetics and phytogeographical structure of Heliosperma (Sileneae, Caryophyllaceae) inferred from chloroplast and nuclear DNA sequences. Molecular Phylogenetics and Evolution 43 (1): 140–155. https://doi.org/10.1016/j.ympev.2006.11.003
Frajman, B., Eggens, F. & Oxelman, B. (2009) Hybrid origins and homoploid reticulate evolution within Heliosperma (Sileneae, Caryophyllaceae)—a multigene phylogenetic approach with relative dating. Systematic Biology 58 (3): 328–345. https://doi.org/10.1093/sysbio/syp030
Greuter, W. (1997) Silene Sect. Heliosperma (Reinchenb.) Ledeb. In: Strid, A. & Tan, K. (Eds.) Flora Hellenica, Vol. 1. Koeltz Scientific Books, Federal Republic of Germany, pp. 293–295.
Grisebach, A. (1843) Spicilegium florae rumelicae et bithynicae exhibens synopsin plantarum, Vol. 1. Prostat apud Fridericum Vieweg et filium, Brunsvigae, 414 pp.
Guangchuang, Y. (2020) Using ggtree to visualize data on tree-like structures. Current Protocols in Bioinformatics 69: e96. https://doi.org/10.1002/cpbi.96
Halbritter, H. & Svojtka, N. (2016) Heliosperma alpestre. In: PalDat - A palynological database. Available from: https://www.paldat.org/pub/Heliosperma_alpestre/301630 (accessed 10 March 2022).
Hoseini, E., Ghahremaninejad, F., Assadi, M. & Nejati Edalatiyan, M. (2017) Seed micromorphology and its implication in subgeneric classification of Silene (Caryophyllaceae, Sileneae). Flora 228: 31–38. https://doi.org/10.1016/j.flora.2017.01.006
Husson, F., Josse, J. & Pagès, J. (2010) Principal component methods-hierarchical clustering – partitional clustering: why would we need to choose for visualizing data? Technical Report. [17 pp.]
Jacquin, J.N. (1773) Florae Austriacae, sive, Plantarum selectarum in Austriae archiducatu: sponte crescentium icones, ad vivum coloratae, et descriptionibus, ac synonymis illustratae, Vol. 1. typis Leopoldi Joannis Kaliwoda, aulae imperialis typographi, Viena, 61 pp. + 100 tabs. https://doi.org/10.5962/bhl.title.457
Kassambara, A. & Mundt, F. (2020) Factoextra: extract and visualize the results of multivariate data analyses. R package version 1.0.7.
Lê, S., Josse, J. & Husson, F. (2008) FactoMineR: an R package for multivariate analysis. Journal of Statistical Software 25: 1–18. https://doi.org/10.18637/jss.v025.i01
Linneaus, C. (1973) Species plantarum, Vol. 1. Salvius, Stockholm, 560 pp.
Malý, K. (1903) Heliosperma (Silene) Retzdorffianum. Österreichische Botanische Zeitschrift 53: 357–359. https://doi.org/10.1007/BF01794437
Malý, K. (1907) Bemerkungen Uber die Arten der Gattung Heliosperma aus der Verwandtschaft des H. Retzdorffianum. In: Hoernes, M. (Eds.) Wissenschaftliche Mitteilungen aus Bosnien und der Herzegovina, Vol. 10. Adolf Holzhausen, Vienna, pp. 629–634.
Markgraf, F. (1931) Pflanzen aus Albanien 1928. Denkschriften der Kaiserlichen Akademie der Wissenschaften Klasse 102 (8): 317–360.
Melzheimer, V. & Polatschek, A. (1992) Revision des Silene (Heliosperma) pusilla agg. (Caryophyllaceae) für die Flora Österreichs. Phyton (Horn, Austria) 31: 281–306.
Mouchet, M., Guilhaumon, F., Villéger, S., Mason, N.W.H., Tomasini, J.-A. & Mouillot, D. (2008) Towards a consensus for calculating dendrogram-based functional diversity indices. Oikos 117 (5): 794–800. https://doi.org/10.1111/j.0030-1299.2008.16594.x
Neumayer, H. (1923) Einige Fragen der speziellen Systematik, erläutert an einer Gruppe der Gattung Silene. Oesterreichische Botanische Zeitschrift 72: 276–287. https://doi.org/10.1007/bf01660057
Niketi?, M. & Stevanovi?, V. (2007) A new species of Heliosperma (Caryophyllaceae) from Serbia and Montenegro. Botanical Journal of the Linnean Society 154: 55–63. https://doi.org/10.1111/j.1095-8339.2007.00643.x
Niketi?, M., Stevanovi?, V. & Tomovi?, G. (2007) Nomenclatural and taxonomic notes on the flora of Serbia and Balkan Peninsula. I. Caryophyllaceae. Archive of Biological Science 59 (4): 387–396. https://doi.org/10.2298/ABS0704387N
Pagès, J. (2004) Analyse Factorielle de Donnees Mixtes. Revue Statistique Appliquee 4: 93–111.
Pan?i?, J. (1875) Elenchus plantarum vascularum quas aestate a. 1873. in Crna Gora. Legit Dr. J. Pan?i?. In typgraphia status, Belgradi, 106 pp. https://doi.org/10.5962/bhl.title.9849
Petchey, O.L. & Gaston, K.J. (2002) Functional diversity (FD), species richness and community composition. Ecology Letters 5 (3): 402–411. https://doi.org/10.1046/j.1461-0248.2002.00339.x
POWO (2022) Plants of the World Online. Available from: http://www.plantsoftheworldonline.org/ (Accessed 30 May 2022).
R Core Team (2021) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: https://www.R-project.org/ (Accessed 10 March 2022).
Rafinesque, C.S. (1840) Autokon Botanikon, Or, Botanical Illustrations of 2500 New, Rare Or Beautiful Trees, Shrubs, Plants, Vines, Lilies, Grasses, Ferns &c, of All Regions, But Chiefly North America, with Descriptions &c and 2500 Self Figures Or Specimens. Arnold Arboretum, Philadelphia, 200 pp.
Reichenbach, H.G.L. (1841) Der Deutsche Botaniker Erster Band. Das Herbarienbuch: Erklärung des natürlichen Pflanzensystems, systematische Aufzählung, Synonymik und Register der bis jetzt bekannten Pflanzengattungen. Der Arnolbitchen Buchhanblung, Dresden, Leipzig, 240 pp.
Reichenbach, H.G.L. (1844) Icones florae Germanicae et Helveticae, simul Pedemontanae, Tirolensis, Istriacae, Dalmaticae, Austriacae, Hungaricae, Transylvanicae, Moravicae, Borussicae, Holsaticae, Belgicae, Hollandicae, ergo Mediae Europae. Friedrich Hofmeister, Leipzig, 84 pp.
Schliep, K., Potts, A.J., Morrison, D.A. & Grimm, G.W. (2017) Intertwining phylogenetic trees and networks. Methods in Ecology and Evolution 8: 1212–1220. https://doi.org/10.7287/peerj.preprints.2054
Sheidai, M., Nikoo, M. & Gholoipour, A. (2008) Cytogenetic variability and new chromosome number reports in Silene L. species (Sect. Lasiostemones, Caryophyllaceae). Acta Biologica Szegediensis 52 (2): 313–319.
Shuka, D., Tan, K., Hallaçi, B. & Shuka, L. (2020) Additions to the flora of North Albania. Phytologia Balcanica 26 (3): 517–522. https://doi.org/10.3372/wi.38.38206
Singh, R.J. (2003) Plant Cytogenetics, 2<sup>nd</sup> ed. CRC Press, Boca Raton, 512 pp.
Thiers, B. (2022 [continuously updated]) Index herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden’s virtual herbarium, New York. Available from: http://sweetgum.nybg.org/science/ih/ (Accessed 10 March 2022).
Trucchi, E., Frajman, B., Haverkamp, T.H.A., Schönswetter, P. & Paun, O. (2017) Genomic analyses suggest parallel ecological divergence in Heliosperma pusillum (Caryophyllaceae). New Phytologist 216: 267–278. https://doi.org/10.1101/044354
Visiani, R. de & Pan?i?, J. (1864) Plantae serbicae rariores aut novae. Memorie dell’Imperial Regio Istituto Veneto di Scienze, Lettere ed Arti 12: 461–478.
Waldstein, F.A. von & Kitaibel, P. (1812) Descriptiones et icones plantarum rariorum Hungariae. Vol. 3. Typis Matthiae Andreae Schmidt, Wien, 88 pp.
Wang, Y., Wang, J., Lai, L., Jiang, L., Zhuang, P., Zhang, L., Zheng, Y., Baskin, J.M. & Baskin, C. (2014) Geographic variation in seed traits within and among forty-two species of Rhododendron (Ericaceae) on the Tibetan plateau: Relationships with elevation, habitat, plant height, and phylogeny. Ecology and Evolution 4 (10): 1913–1923. https://doi.org/10.1002/ece3.1067
Wickham, H., Averick, M., Bryan, J., Chang, W., McGowan, L.D, François, R., Grolemund, G., Hayes, A., Henry, L., Hester, J., Kuhn, M., Pedersen, L.T., Miller, E., Bache, S.M., Müller, K., Ooms, J., Robinson, D., Seidel, D.P., Spinu, V., Takahashi, K., Vaughan, D., Wilke, C., Woo, K. & Yutani, H. (2019) Welcome to the tidyverse. Journal of Open Source Software 4 (43): 1686. https://doi.org/10.21105/joss.01686