G Model PPEES-25121; No. of Pages 12 ARTICLE IN PRESS Perspectives in Plant Ecology, Evolution and Systematics xxx (2011) xxx–xxx Contents lists available at ScienceDirect Perspectives in Plant Ecology, Evolution and Systematics journal homepage: www.elsevier.de/ppees Biological Flora of Central Europe Biological flora of Central Europe: Aster amellus L. (Asteraceae) Zuzana Münzbergová a,b,∗, Jana Raabová c, Sílvia Castro a,d, Hana Pánková a a Department of Botany, Faculty of Science, Charles University in Prague, Benátská 2, 128 01 Prague, Czech Republic b Institute of Botany, Academy of Sciences, Zámek 1, 252 43 Pruhonice,˚ Czech Republic c Department of Botany, National Museum, Cirkusová 1740, 193 00 Praha 9, Czech Republic d CFE, Centre for Functional Ecology, Department of Life Sciences, University of Coimbra, PO Box 3046, 3001-401 Coimbra, Portugal article info abstract Article history: Aster amellus L. (Asteraceae) is a polymorphic aggregate of taxa. The species aggregate is distributed Received 18 May 2010 through Central and Eastern Europe and Western Asia. The habitats of the species include grasslands, Received in revised form 1 February 2011 clearings, edges, slopes, waysides and open forests. Typical habitats of this species have declined over the Accepted 14 March 2011 last decades, and A. amellus became endangered in many parts of Central Europe. This paper deals with taxonomy, morphology, distribution, habitat requirements, life cycle and biology of this species. Special Keywords: emphasis is given to the differences between diploid and hexaploid plants, which co-occur in Central Central Europe Europe. Ecology Flow cytometry © 2011 Elsevier GmbH. All rights reserved. Genetics Polyploidy Species biology Taxonomy and morphology A. amellus is the type species for the genus Aster, family Aster- aceae and order Asterales (Jones, 1980; Pennisi, 2001). The name Taxonomy Aster comes from the Ancient Greek word astér, which means “star” and refers to the shape of the flower head. Aster s.l. is a mor- Aster amellus L. Sp. Pl. 873, 1753 (Asteraceae) – Italian aster phologically heterogeneous and geographically widespread genus, Heterotypic synonyms: comprising over 400 species, with centres of diversity in North Aster amelloides Hoffm. Gött. Anz. Gel. Sachen 20: 1325, 1800. America and Eurasia. The genus has a complex taxonomic history, – Aster ottomanus Velen. Sitzungsber. Königl. Böhm. Ges. Wiss., with several partial taxonomic treatments attempting to iden- Math.-Naturwiss. Cl., 1890: 48, 1890. tify natural groups (reviewed in Jones, 1980; Jones and Hiepko, Aster bessarabicus Rchb. Fl. Germ. Excurs. 246, 1831–1832. – 1981; Nesom, 1994a; Xiang and Semple, 1996). Recent morpho- Aster amellus subsp. bessarabicus (Rchb.) Soó Acta Bot. Acad. Sci. logic and molecular studies revealed that the genus is generally Hung. 12: 366, 1966. – Aster scepusiensis Kitaibel ex Kanitz Linnaea restricted to the Old World species (Aster s.s. with about 180 32: 373, 1863. – Aster amellus subsp. scepusiensis (Kanitz) Dostál species; Nesom, 1994a,b; Noyes and Rieseberg, 1999), and most of Folia Mus. Rerum Natur. Bohem. Occid., Bot., 21: 12, 1984. – Aster the North American asters belong to other related genera (Nesom, amelloides Besser Enum. Pl. 33, 1821 [non Hoffm. 1800], homonym. 1994b; Xiang and Semple, 1996; Noyes and Rieseberg, 1999). Phy- Aster ibericus M. Bieb. Fl. Taur.-Caucas. 2: 311, 1808. – Aster amel- logenetic studies based on nucleotide sequence data from the lus subsp. ibericus (M. Bieb.) V.E. Avet. Biol. Zurn.ˇ Armenii 25(10): internal transcribed spacers (ITS) of nuclear ribosomal DNA and 63, 1972. amplified fragment length polymorphism (AFLP) markers suggest Aster hirtus K. Koch Linnaea 23: 701, 1851 [non Scop. 1772]. – that Old World Aster is a polyphyletic group, with independent lin- Aster elegans Willd. Sp. Pl., ed. 4 [Willd.] 3(3): 2042, 1803. – Amellus eages derived from possibly distant southern hemisphere ancestors officinalis Gaterau Fl. Montauban 147, 1789, nom. illeg. – Amellus (Noyes and Rieseberg, 1999; Cammareri et al., 2004). vulgaris Opiz Sezn. Rostl. Kvet.ˇ Ces.ˇ 14, 1852, nom. illeg. Phylogenetic and cladistic studies performed so far within the genus Aster reveal a close relationship between A. amellus and A. alpinus (AFLP analysis, Cammareri et al., 2004; cladistic analysis, Jones and Young, 1983; Nesom, 1994a). Xiang and Semple (1996) ∗ Corresponding author at: Department of Botany, Faculty of Science, Charles Uni- found different phylogenetic relationships within Aster based on versity in Prague, Benátská 2, 128 01 Prague, Czech Republic. Tel.: +420 221 951 636; divergence of cpDNA. This classification is, however, suspicious as fax: +420 221 951 645. E-mail address: [email protected] (Z. Münzbergová). the sample of A. amellus used in the study of Xiang and Semple 1433-8319/$ – see front matter © 2011 Elsevier GmbH. All rights reserved. doi:10.1016/j.ppees.2011.03.002 Please cite this article in press as: Münzbergová, Z., et al., Biological flora of Central Europe: Aster amellus L. (Asteraceae). Perspect. Plant Ecol. Evol. Syst. (2011), doi:10.1016/j.ppees.2011.03.002 G Model PPEES-25121; No. of Pages 12 ARTICLE IN PRESS 2 Z. Münzbergová et al. / Perspectives in Plant Ecology, Evolution and Systematics xxx (2011) xxx–xxx (1996) was an octoploid plant from a garden in Quebec, Canada and els cannot be morphologically distinguished. A molecular approach it may in fact be a hybrid of A. amellus with some Eurybia species. also revealed that the two cytotypes cannot be distinguished based A. amellus (Fig. 1) is a polymorphic aggregate of taxa, includ- on isozyme profiles, thus sharing a common gene pool (Mandáková ing three cytotypes: diploids (2n =2x = 18), tetraploids (2n =4x = 36) and Münzbergová, 2008). The two cytotypes are also not separated and hexaploids (2n =6x = 54) (Merxmüller et al., 1976; Májovsky,´ based on cpDNA (Castro et al., unpubl.). Castro et al. (unpubl.) dis- 1978). The aggregate of the species has received several taxonomic tinguished until now four major clades in Central Europe, where treatments based on both morphological and caryological charac- two clades contained both diploid and hexaploid individuals, sug- ters. Tamamshyan (1990) and Májovsky´ (1978) distinguished three gesting multiple independent origin of hexaploids. These results independent taxa within the aggregate: A. amellus L., A. ibericus support the treatment of Aster amellus as one polymorphic species Stev. and A. amelloides Bess. non Hoffm. instead of multiple independent taxa. According to Májovsky´ (1978) the classification of the species Hegi (1979) indicates that the species is very variable in its leaf as three independent taxa is justified by their morphological dif- shape and hairs without any needs to distinguish new taxa. Some ferences, different chromosome numbers, distinct expected areas plants with reddish edge of involucral bracts were described as f. of origin and different evolutionary histories. Following Májovsky´ lauticeps Beck and similarly plants with strongly pubescent leaves (1978), A. amellus L. includes diploid plants with small heads and were described as var. hispidus DC. However, the taxonomic value oval obtuse involucral bracts, distributed from sub-Mediterranean of these forms is doubtful (Hegi, 1979). to temperate sub-continental Europe and to western Siberia. A. Due to the unclear taxonomical treatment of this group and ibericus Stev. includes tetraploid plants with big heads, long disk- to the recent morphological and molecular studies (Mandáková florets and lanceolate acute hairy involucral bracts, distributed and Münzbergová, 2008), we use a conservative approach, fol- from sub-Mediterranean mountains to sub-temperate western lowed also in Flora Europaea (Merxmüller et al., 1976), and consider Asia. A. amelloides Bess. non Hoffm. includes hexaploid plants with A. amellus as a polymorphic species, including all the cytotypes, big heads and lanceolate acute glabrous involucral bracts, dis- through the paper. Lot of biological data on A. amellus is available tributed from sub-Mediterranean via sub-temperate to eastern only from limited geographic area of this species, not including sub-continental Europe and to western Asia. all ploidy levels and subspecies. Because we are aware of the lim- Contrary to Májovsky´ (1978), studies on chromosome counts itations of these findings for the whole aggregate of the taxa, we of these species (Huziwara, 1962; Magulaev, 1986) report that A. always report the geographic area of the specific studies. For exam- amellus L. are diploid plants, and A. ibericus Stev. and A. amelloides ple, comparisons of habitats, life cycle, phenology and reproduction Bess. non Hoffm. are both hexaploid plants. between diploid and hexaploid plants come mainly from our own Tamamshyan (1990) does not report the ploidy level of the research on A. amellus in the Czech Republic. three taxa, but distinguishes the same three taxa as Májovsky´ (1978) based on plant morphology. According to Tamamshyan Morphology (1990), A. amellus plants are glabrous or subglabrous. The outer bracts of involucre are subglabrous. Inner bracts are lanceolate Description of plant morphology is given for the whole polymor- and both types of bracts are coloured. Lower leaves are elliptic- phic A. amellus species. Where the information is available, we also spathulate and the stem is often reddish. In contrast, A. amelloides provide separate information for diploid and hexaploid cytotypes and A. ibericus