Environmental Management (2017) 59:966–981 DOI 10.1007/s00267-017-0833-4 Specific Vicariance of Two Primeval Lowland Forest Lichen Indicators 1 2 Dariusz Kubiak ● Piotr Osyczka Received: 6 September 2016 / Accepted: 3 February 2017 / Published online: 15 February 2017 © The Author(s) 2017; This article is published with open access at Springerlink.com Abstract To date, the lichens Chrysothrix candelaris and and it requires forests with high proportion of deciduous Varicellaria hemisphaerica have been classified as accurate trees in a wide landscape scale. Local landscape-level primeval lowland forest indicators. Both inhabit particularly habitat continuity is more important for this species than the valuable remnants of oak-hornbeam forests in Europe, but current age of forest stand. Regardless of the indicative tend toward a specific kind of vicariance on a local scale. value, localities of both lichens within oak-hornbeam for- The present study was undertaken to determine habitat ests deserve the special protection status since they form factors responsible for this phenomenon and verify the unique assemblages of exclusive epiphytes, including those indicative and conservation value of these lichens. The main with high conservation value. spatial and climatic parameters that, along with forest structure, potentially affect their distribution patterns and Keywords Oak-hornbeam forest ● Lichen biota ● Lichen abundance were analysed in four complexes with typical conservation ● Environment evaluation ● Host-tree ● Habitat oak-hornbeam stands in NE Poland. Fifty plots of 400 m2 requirements each were chosen for detailed examination of stand struc- ture and epiphytic lichens directly associated with the indicators. The study showed that the localities of the two species barely overlap within the same forest community in a relatively small geographical area. The occurrence of Introduction Chrysothrix candelaris depends basically only on micro- habitat space provided by old oaks and its role as an indi- The spared remnants of natural deciduous or mixed forests cator of the ecological continuity of habitat is limited. in Europe represent a typical zonal formation constituting Varicellaria hemisphaerica is not tree specific but a suffi- the dominant type of potential vegetation over large areas of ciently high moisture of habitat is essential for the species the continent (Ellenberg 1988). Contrary to commercial forests, used primarily as a source of wood, deciduous forests have high environmental value, as they are char- acterised by great diversity of tree species and constitute Electronic supplementary material The online version of this article important refuges for various organisms (Faliński and (doi:10.1007/s00267-017-0833-4) contains supplementary material, Mułenko 1995; Barbier et al. 2008). which is available to authorized users. Lichens are a conspicuous component, occupying the * Dariusz Kubiak entire space of forest communities (Coxson and Nadkarni [email protected] 1995; Sillett and Antoine 2004; Seaward 2008; Ellis 2012). While they occur in various ecological groups, epiphytes 1 Department of Mycology, University of Warmia and Mazury in Olsztyn, Oczapowskiego 1A, 10-719 Olsztyn, Poland clearly dominate in deciduous and mixed forests of the 2 temperate zone. Many lichens are closely related to a spe- Department of Polar Research and Documentation, Institute of fi Botany, Jagiellonian University, Kopernika 27, 31-501 Kraków, ci c habitat and thus may possess indicative value (e.g., Poland Nimis and Martellos 2001). Both external and internal Environmental Management (2017) 59:966–981 967 factors, e.g., climate, landscape form, composition and preserved European lowland forests is a mystery. It is dif- maturity of trees, and biological interactions (Leppik and ficult to find a study on lichens of old-growth lowland forest Jüriado 2008; Moning et al. 2009; Marini et al. 2011; Hauck which does not report the presence of one of these species, et al. 2013; Nascimbene et al. 2013), affect lichen diversity yet the simultaneous existence of both populations in in forest complexes. The relative effect of these factors on similar abundance within a single forest complex is lichen biota development is difficult to assess (Pinho et al. observed only sporadically (Cieśliński and Tobolewski 2008; Giordani and Brunialti 2015). Moreover, local cli- 1988; Zalewska 2012; Malíček and Palice 2013; Kubiak matic fluctuations, elevation, and/or land-use intensity drive et al. 2016). Therefore, our research was aimed at answering the local specificity of lichen diversity (Loppi et al. 1997; the following questions: (1) Does the current structure of a Giordani 2006; Wolseley et al. 2006; Giordani and Incerti forest stand affect the distribution of the examined species? 2008). Generally, old-growth, least-affected lowland If so, to what extent? (2) If so, what kinds of forest stands deciduous forests are characterised by a high level of lichen constitute optimal habitats for these species? (3) If not, are diversity, and many rare species are associated with old there other factors responsible for their distribution pat- trees (Hyvärinen et al. 1992; Price and Hochachka 2001; terns? (4) Are specific assemblages of lichens directly Fritz et al. 2008a, b; Ranius et al. 2008a; Nascimbene et al. associated with these species due to similar habitat 2010a; Bartels and Chen 2012, 2014). requirements? Numerous endangered epiphytic lichens are stenotopic These questions seem particularly crucial in the context and specially adapted to certain habitat types (Bartels and of the practical use of these lichens as environmental indi- Chen 2015). These species may be a signal determinant of cators. Moreover, we hypothesise that: (1) the distribution the natural condition of the forest environment (McCune patterns of the examined species are not accidental and 2000). The use of a limited number of bioindicators is an result from their narrow ecological amplitude; (2) their alternative and practical approach in environmental assess- vicariance is caused by differences in forest stand structure; ments (Nitare 2000). The great advantages of such an (3) both lichen indicators are leading representatives of approach are a short study period, simplicity, and low cost unique assemblages of species with similar habitat (Will-Wolf et al. 2002; Gao et al. 2015). However, detailed requirements. data on the distribution and habitat requirements of lichen indicators are required for correct interpretation of field observations and for appropriate conservation actions and Materials and Methods policies (Löhmus and Löhmus 2009; Juriado et al. 2012; Jönsson et al. 2016). Some species may have special con- Study Area servation value, and thus their protection entails that of naturally co-occurring species and entire biocoenoses The study was conducted in the Masurian Lakeland (NE (Nilsson et al. 1995; Roberge and Angelstam 2004; Schei- Poland), almost one-third of which is covered by forests degger and Werth 2009; Ivanowa 2015). Therefore, atten- with a high degree of biodiversity (Faliński 1998). This tion should be also paid to lichen biota directly associated region constitutes a natural corridor constituting an impor- with a particular indicator (Nilsson et al. 1995; Johansson tant environmental communication link in Central Europe and Gustafsson 2001; Nordén et al. 2007; Nascimbene et al. (CORINE biotopes 2016). Field studies were carried out in 2010b). four large forest complexes, in many parts of which oak- The impact of various factors on the local occurrence of hornbeam stands have been preserved in their natural form two epiphytic lichens, Chrysothrix candelaris (L.) J. R. to the present day: the Puszcza Nidzicka f. (abbr. N), Laundon and Varicellaria hemisphaerica (Flörke) I. Puszcza Piska f. (abbr. P), Puszcza Borecka f. (abbr. B), and Schmitt & Lumbsch, within one type of forest community Puszcza Romincka f. (abbr. R) (Fig. 1). The topography of was examined in this study. These species are commonly the region is very diverse, as the landscape was formed used as suitable primeval lowland forest indicators (Coppins during the Pomeranian phase of the last Vistula River gla- and Coppins 2002; Czyżewska and Cieśliński 2003; ciation (ca 15,200 BP) (Kozarski and Nowaczyk 1999). Motiejūnaitė et al. 2004) and are considered typical inha- Denivelations are considerable, reaching as much as bitants of forests of above-average biodiversity (Andersson 120–140 m a.s.l; the highest point is almost 300 m a.s.l. and Kriukelis 2002; Ek et al. 2002). The two species are This region is characterised by zonal climatic variability, morphologically distinctive and easy to identify in the field, with clashes of oceanic, continental, and boreal climate even by non-specialists; therefore they have a wide practical types (Kondracki 1972). Generally, the forest complexes application in field evaluations. Nevertheless, the lack of situated further to the north-east, due to the greater impact parallel contributions of C. candelaris and V. hemi- of arctic air masses and relatively high elevation, experience sphaerica in the epiphytic biota of some of the best- a colder and more humid climate than those in the south- 968 Environmental Management (2017) 59:966–981 Fig. 1 Location of the studied forest complexes in Poland. Total areas of the complexes and general coverage participations of forest stands within the complexes in relation to age (less than 100 years : over 100 years) and kinds of trees (coniferous