Thirty-Year Invasion of the Alien Moss Campylopus Introflexus (Hedw.)
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Biol Invasions https://doi.org/10.1007/s10530-018-1818-9 (0123456789().,-volV)(0123456789().,-volV) INVASION NOTE Thirty-year invasion of the alien moss Campylopus introflexus (Hedw.) Brid. in Poland (East-Central Europe) Jan Zarnowiec_ . Adam Stebel . Damian Chmura Received: 6 March 2018 / Accepted: 19 August 2018 Ó The Author(s) 2018 Abstract Campylopus introflexus is the only inva- found in acidophilous semi-natural forests and inland sive moss listed among the 100 most prominent alien dunes with Corynephorus and Agrostis grasslands, invaders in Europe. The present study shows dot boggy woodland, dry heaths, and ruderal habitats. distribution maps (at 10-years intervals from its Although 30 cryptogam species were found in tufts of introduction in 1986 to the present) and discusses the the alien moss, there is no characteristic species dynamics of its invasion, altitudinal distribution and composition that constantly accompanies its presence. ecological preferences. During the 30 years of its invasion, C. introflexus has spread widely throughout Keywords Adventive moss Á Invasive moss Á Poland and is currently known in 248 locations. A total Cryptogamic plant invasion Á Neophyte Á Secondary of 93.1% of these are below 300 m a.s.l., and the range Á Distribution map maximum altitude is 780 m. With the passage of time since its first appearance this alien moss has occupied larger numbers of substrates and habitats. Currently C. introflexus is a component of 34 plant communities Introduction belonging to 18 habitat types. The most invaded habitats include young forest tree plantations and The invasion of alien species is currently considered to mature managed forest developed from old plantations be among the most important and difficult problems (Pinus sylvestris woodland). This moss is frequently confronting nature protection. Compared with vascu- lar plants, the phenomenon of non-native species invasion occurs on a much smaller scale in mosses. J. Zarnowiec_ (&) Á D. Chmura There is also a much smaller negative impact on Institute of Environmental Protection and Engineering, biodiversity and on socio-economic features (Essl University of Bielsko-Biała, ul. Willowa 2, 43–309 Bielsko-Biała, Poland et al. 2014). On a global scale there are only 106 alien e-mail: [email protected] mosses, including 37 cryptogenic species (Essl et al. D. Chmura 2013). In Europe 21 non-native moss species (1.6% of e-mail: [email protected] the European moss flora) and 11 cryptogenic species (0.8%) have been identified (Essl and Lambdon 2009). A. Stebel In Poland, there are three alien moss species (0.4% of Department of Pharmaceutical Botany, Medical University of Silesia in Katowice, ul. Ostrogo´rska 30, the Polish moss flora), namely Campylopus intro- 41–200 Sosnowiec, Poland flexus, Orthodontium lineare and Leptophascum e-mail: [email protected] 123 J. Zarnowiec_ et al. leptophyllum (Ochyra et al. 2003; Fudali et al. 2009). short or almost lacking. It commonly reproduces The first of these is the most highly invasive and is vegetatively by deciduous leaves and fragile stem tips listed among the 100 most prominent alien invaders which facilitate its short-distance spread. Dioicous, (Lambdon 2009). Molecular studies on Campylopus female shoots have terminal perichaetial buds which introflexus have demonstrated that in its entire native frequently produce sporophytes. Plants are occasion- and secondary ranges it is genetically homogenous ally polysetous; setae yellow–brown to pale brown, (Stech and Dohrmann 2004; Gama et al. 2017). It is 5–15 mm long, curved or sinuose. Capsule brown, thus impossible to determine using genetic methods slightly asymmetric and curved, furrowed and shrun- whether its European invasion is the result of one or ken when dry; spores pale brown, small 10–14 lm; many introductions. Although knowledge of adven- ripe sporophytes have been observed in Poland tive moss invasions remains incomplete, the history of between March and October. C. introflexus spread in Europe is one of the best Native range—C. introflexus is native to the known and documented. These data, however, pri- Southern Hemisphere, where this moss has a pan- marily refer to Western Europe (Klinck 2010 and the temperate range. C. introflexus is widespread in the literature cited there) and to Central Europe (Miku- southern part of South America and Africa, southern la´sˇkova´ et al. 2012;Szucs} et al. 2014), and few data and eastern parts of Australia, in Tasmania and in New refer to East-Central Europe. Zealand. Its range reaches north to New Caledonia, C. introflexus was first discovered in Poland in Re´union, St Helena, while it has been reported from October 1986 (Lisowski and Urban´ski 1989). By 2016 most Subantarctic and some Antarctic islands (Grad- 30 years had passed since the first record, presenting a stein and Sipman 1978;So¨derstro¨m 1992; Ochyra good opportunity to summarize data on the invasion et al. 2008; Klinck 2010). and ecological preferences of this moss in Poland. Non-native, secondary range—Outside its native Moreover, production of a distribution map in this range it was first recorded in 1941 at Washington, country was attempted nearly 10 years ago (Fudali West Sussex in Great Britain and in 1942 at Howth et al. 2009). The present study aims to answer the near Dublin in Ireland (Richards 1963). On mainland following main questions: (i) what is the current Europe, it was initially observed in 1954 in Brittany in distribution of C. introflexus in Poland and what was France (Størmer 1958) before spreading rapidly the dynamic of its spread? (ii) What substrates and throughout Europe (Gradstein and Sipman 1978; habitats were invaded by C. introflexus and were its So¨derstro¨m 1992; Smith 2004; Lambdon 2009; Klink ecological preferences changed as a result? (iii) With 2010). The current non-native range of C. introflexus which native species of bryophytes and lichens does C. in Europe extends north to Iceland (Klinck 2010), introflexus coexist? south-west Norway (Øvstedal 1978) and Sweden (Ha¨llingba¨ck et al. 1985); north-east—along the Baltic coasts—by the Kaliningrad Province of Russia Methods (Razgulyaeva et al. 2001), Lithuania (Jukoniene_ et al. 2015), Latvia (Abolina and Reriha 2004)to The study species Estonia (Vellak et al. 2009); east to Ukraine (Lobachevska and Sokhanchak 2010), Hungary Campylopus introflexus is a perennial, acrocarpous, (Szucs} et al. 2014), Turkey (Ros et al. 2013); south medium-sized moss that forms yellowish to olive to Italy and the Mediterranean islands (Bealarics, green or rarely dark green loose tufts or dense mats; Corsica, Sardinia) (Ros et al. 2013), and west to hoary when dry. Shoots are 0.5–7.0 cm tall, with Portugal (Se´rgio 1979) it has also been reported from slender internodes and swollen nodes, tomentum the Azores, Madeira, and the Canary Islands in reddish-brown present or almost absent. Leaves Macaronesia (Ros et al. 2013). C. introflexus fre- 2.5–6.0 mm in length, straight, erecto-patent when quently grows along the Pacific coasts of North moist, appressed when dry, lanceolate with entire America and has also been recorded in California, margins. Costa strong, excurrent in a dentate hyaline Oregon, Washington, and southern British Columbia hair-point, which is conspicuously 90° reflexed; (Frahm 2007; Carter 2014). rarely, in plants growing in shade, hyaline hair-tip 123 Thirty-year invasion of the alien moss Campylopus introflexus (Hedw.) Brid. in Poland… Ecology—The habitats that C. introflexus occupies recognized, among other attributes, by the conspicu- in its native range are similar to those in the secondary ously 90° reflexed hair points and is unlikely to be (alien) range. It has a high ecological tolerance, and its confused with any other species in Poland. The revised populations are associated with both undisturbed and specimens and published data on its occurrence refer anthropogenic sites. In Europe C. introflexus occurs on to 248 localities, including collections from 91 stands, a wide range of substrata and in greatly varied habitats. in which we made complete observations and pro- It is seen most frequently on acid sandy and sandy- vided descriptions of the ecological situation. In this gravelly soils, bare dry peat and less often on decaying group there are 60 unpublished stations. For 113 sites, rotten wood, rocks, waste deposits, and roofs of information on species co-occurring with C. intro- buildings. Its habitats include coastal grey dunes, dry flexus is available. All information on habitat, altitude, sandy grassland vegetation, moorland, raised and substrates, and associated species was included in a blanket bogs (mostly disturbed or burned), coniferous database. The data gathered during fieldwork and forest, forestry tracks, plantations, forest ridges and information from herbarium labels were used to clearings and margins of fish ponds (Richards and describe the ecological preferences of the Polish Smith 1975; Smith 2004; Lambdon 2009; Klinck populations of C. introflexus. A herbarium-based 2010; Mikula´sˇkova´ et al. 2012). In North America C. method has previously been successfully used, among introflexus primarily colonizes poor acid soils along others, to assess changes in the frequency of mosses in the Pacific coasts in Pine forest and on stabilized sand Sweden (Hedena¨s et al. 2002) and to reconstruct the dunes. It thrives on sandy soils along trails, at the base spread of invasive plants (Delisle et al. 2003). of trees, on the roofs of buildings, and on peat in bogs The distribution of C. introflexus is shown on dot (Frahm 2007; Carter 2014). In active volcanic zones in maps at 10-years intervals (Fig. 1a–c). We also Antarctica, Iceland, and in central Italy, this moss prepared a curve showing the spread of this alien grows on geothermal ground and around active vents species in Poland (Fig. 1e). Its distribution pattern is emitting sulphur and steam (Chiarucci et al.