Adenophora Liliifolia: Condition of Its Populations in Central Europe
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ACTA BIOLOGICA CRACOVIENSIA Series Botanica 58/2: 83–105, 2016 DOI: 10.1515/abcsb-2016-0018 ADENOPHORA LILIIFOLIA: CONDITION OF ITS POPULATIONS IN CENTRAL EUROPE ROMANA PRAUSOVÁ1a*, LUCIE MAREČKOVÁ2a, ADAM KAPLER3, L’UBOŠ MAJESKÝ2, TÜNDE FARKAS4, ADRIAN INDREICA5, LENKA ŠAFÁŘOVÁ6 AND MILOSLAV KITNER2 1University of Hradec Králové, Faculty of Science, Department of Biology, 500 02 Hradec Králové, Czech Republic 2Palacký University in Olomouc, Faculty of Science, Department of Botany, Šlechtitelů 27, 783 71 Olomouc-Holice, Czech Republic 3PAS Botanical Garden – Center for Biological Diversity Conservation in Powsin, Prawdziwka 2, 02-973 Warsaw 76, Poland 4Aggteleki Nemzeti Park Igazgatóság, Tengerszem oldal 1, 3759 Jósvafő, Hungary 5Transilvania University of Brasov, Faculty of Forestry, Şirul Beethoven – 1, 500123 Braşov, Romania 6East Bohemian Museum in Pardubice, Zámek 2, 530 02 Pardubice, Czech Republic Received June 16, 2016; revision accepted September 30, 2016 This study deals with populations of the European-South-Siberian geoelement Adenophora liliifolia (L.) A. DC. in the Czech Republic, Slovakia, Hungary, Romania, and Poland, where this species has its European periphery distri- bution. We studied the population size, genetic variability, site conditions, and vegetation units in which A. liliifolia grows.Keywords: Recent and historical localities of A. liliifolia were ranked into six vegetation units of both forest and non-for- est character. A phytosociological survey showed differences in the species composition among localities. Only a weak pattern of population structure was observed (only 22% of total genetic variation present at the interpopulation level, AMOVA analysis), with moderate values for gene diversity (Hj = 0.141) and polymorphism (P = 27.6%). Neighbor- joining and Bayesian clusterings suggest a similar genetic background for most of the populations from Slovakia, the Czech Republic, and Poland, contrary to the populations from Hungary, Romania, as well as two populations from Central and South Slovakia. This might be explained by a relatively recent fragmentation of the A. liliifolia popula- tions in Central Europe. Nevertheless, it seems that several populations in Romania, South Hungary, and Slovakia were isolated for a longer period of time and their genetic differentiation is more evident. Keywords: AFLP, Campanulaceae, European periphery distribution, declining population, European- South-Siberian geoelement, genetic variability, vegetation INTRODUCTION (Michx.) Kunth ex Iljinsk. (Denk et al., 2001)), or in the Balkan Peninsula (e.g., Picea omorika The present-day flora of Central Europe reflects (Pančić) Purk. (Ravazzi, 2002)). Some species, its geographic position, varied geology and topog- such as Ligularia sibirica (L.) Cass. (Šmídová et raphy, as well as climate and vegetation history, al., 2011) or Pedicularis sudetica Willd. (Hendrych and it is influenced by glacial cycles during the and Hendrychová, 1988), became glacial or postgla- Quaternary Period (Grulich, 2012; Kaplan, 2012). cial relicts, but also new local endemics appeared, Numerous species that are extinct in Central e.g., Galium sudeticum Tausch, G. cracoviense Europe survived in Eastern Asia (e.g., Platycladus Ehrend., Cochlearia polonica E. Froelich (Cieślak orientalis (L.) Franco (Farjon and Filer, 2013)), et al., 2007, 2010, 2015; Cieślak and Szeląg, 2009, in Transcaucasia (e.g., Pterocarya pterocarpa 2010; Kolář et al., 2013, 2015) or Sorbus sudet- ica (Tausch) Bluff et al. (Kaplan, 2012). Climatic * Corresponding author, email: [email protected] a These two authors contributed equally to this work. PL ISSN 0001-5296 © Polish Academy of Sciences and Jagiellonian University, Cracow 2016 84 Prausová et al. changes during the Quaternary Period strongly considered as a species of least concern (Bilz et al., affected the species composition and the species 2011); and it is threatened by vigorous shrubby distribution in Europe (Szafer, 1946–47, 1954; vegetation and by inappropriate forest manage- Kaplan, 2012). Since the Neolithic Period, humans ment (Anonymous, 2009). A. liliifolia is considered have become another important factor affecting the a plant species of European Community interest, regional floras (Szymura, 2012; Hejcman et al., whose conservation requires designation of special 2013; Roleček et al., 2014; Plieninger et al., 2015). areas of conservation. Moreover, it is an indicator Many species, such as A. liliifolia (L.) A. DC, species of thermophilous forest hotspots, signaling changed their distribution and their recent pres- remnant pools of biodiversity (Kiedrzyński et al., ence in floras is highly influenced by human activity. 2015). A typical habitat of A. liliifolia is the cop- A. liliifolia is considered to represent the European- pice, which is a formerly widespread way of forest South-Siberian geoelement, which tolerates extreme management. However, changes in landscape man- continental climate with a short growing season, agement during the last two centuries caused the warm but short summers, and long winters with extinction of this species because of the shady and severe frosts (Kucharczyk, 2007; Kaplan, 2012; more eutrophicated high forests (often with coni- Kucharczyk et al., 2014). The centre of A. liliifolia fers) that replaced the coppices (Szymura, 2012; distribution is in Western Asia-Southern Siberia, Müllerová et al., 2015). Today A. liliifolia grows in and from there it extends to Mongolia and Western lowlands in small populations in remnants of for- China in the East, and to the North-West of Turkey, mer light oak forests, their ecotones, and adjacent and to South-, Eastern-, Central- Europe up to meadows. At higher altitudes it grows on the rocky Western Europe in the West (Tacik, 1971; Fedorov, outcrops in beech forests (Moser, 1999; Dražil, 1978; Deyuan et al., 2011; Urgamal, 2014). 2002), and in the portions of riparian forests Although Smelansky et al. (2004) reported A. liliifo- receiving large quantites of sunlight (Siklósi, 1984; lia as a common species in the steppes and forest- Farkas and Vojtkó, 2012, 2013). These ecological steppes in Southern Siberia, Boronnikova (2009) demands make A. liliifolia a suitable model species reported a 25% decrease of populations in the Perm for studying the changes and the impact of human region (Ural, Russia) during the last 15 years due activities on populations of species with similar to agricultural activities in the territory. Also, in the characteristics. whole Central European region, A. liliifolia popula- In spite of the critical conservation status of tions are declining not only in the number of locali- A. liliifolia in Europe, no large-scale population ties, but also in the number of plants representing genetic studies have been done so far. Only two a single population. studies have investigated the population struc- A. liliifolia is scattered across Europe and ture of A. liliifolia: Boronikova (2009) analyzed forms isolated populations in Germany (Meusel four populations from the Ural region (Perm, and Jäger, 1992; Castroviejo et al., 2010), Russia), and Manole et al. (2015) described the Austria, Switzerland (Moser, 1999), Italy, Czechia genetic diversity of one A. liliifolia population (Martinovský, 1967; Kovanda, 2000), Poland from Romania. The present study aimed to pro- (Witkowski et al., 2003; Korzeniak and Nobis, vide overall information on the current condition 2004; Ciosek, 2006; Kapler et al., 2015), Slovakia of A. liliifolia populations in Central Europe, and (Goliášová and Šípošová, 2008), Hungary (Farkas thus to better know the factors threatening this spe- and Vojtkó, 2012, 2013; Vojtkó, 2013), Croatia, cies and suggest appropriate management for the Bosnia and Herzegovina, Montenegro, Serbia current populations. We performed: 1) a survey and (Vladimirov et al., 2009; Vukojičić et al., 2011), comparison of A. liliifolia populations in Czechia, Romania (Jones et al., 2010; Indreica, 2011), and Slovakia, Hungary, Romania, and Poland; as well as Slovenia (Babij, 2004; Acetto, 2007). In Belarus, 2) a screening of the genetic variability and relation- the species was thought to be extinct (Kozlovskaja, ship among the studied populations by means of 1978), but one population at Sporowski Zakaznik Amplified Fragment Length Polymorphism (AFLP). was restored with plants multiplied in vitro and cultivated in the Minsk Botanical Garden of the Belarussian Academy of Sciences (Wiliams and MATERIALS AND METHODS Gotin, 2012). The information about A. liliifolia from France (Schnittler and Günther, 1999) and STUDY SPECIES Bulgaria (Dimitrov, 2002) is uncertain, as no her- barium records from France and the current The lilyleaf ladybells Adenophora liliifolia is a her- Bulgarian territory exist. baceous perennial diploid (2n = 34) plant from A. liliifolia is protected in Europe according to the Campanulaceae family with erect, leafed and the Directive on the conservation of natural habi- branched stems. The root is spindle-shaped or tats and of wild fauna and flora (92/43/EEC); it is branched. The plant usually grows to a height of Adenophora liliifolia in Central Europe 85 40–90 cm (Kovanda, 2000), although plants with in all the countries. The condition of the locality heights of 205 cm have been observed in Poland and its changes were observed during the moni- (Ciosek, 2006). Basal leaves and leaves of young toring of populations in 2012–2013. The danger plants are long petiolate, with cordate to rounded of possible damage