Tree species diversity change in natural regeneration of a beech forest under different management
Introduction
Temperate forests have been significantly altered by land use (Burrascano et al., 2013), even more so than other forest biomes (Silander, 2001, Terek and Dobrovič, 2013), because they are distributed in the mid-latitudes were environmental factors such as climate are optimal to support human population. Land-use is considered to have an impact on biodiversity (Ódor et al., 2014), but the extent and consequences of different management intensities are not yet known (Fischer et al., 2010). Biodiversity plays an important role in decision making and is recognised as a key element of sustainable forest management world-wide (FAO, 2006, Baycheva et al., 2013). Well-targeted forest biodiversity conservation policies will become even more strategic in the future (Barbati et al., 2014). Therefore, knowledge on whether or not, and to what extent the management strategies influence future species biodiversity is of high importance.
Beech is the most abundant broadleaved tree species in Europe (e.g. Ellenberg, 1996) and plays an important economic (Taghiyari, 2013, Shahverdi et al., 2013) and ecological (Drobyshev et al., 2014, Hlásny et al., 2014) role in sustainable forest management. European beech (Fagus sylvatica L.) also dominates the potential natural vegetation in a large part of Central Europe (Bohn et al., 2004). It is characterised as a late-successional, shade-tolerant species with a broad ecological tolerance (Ellenberg, 1996) and good regeneration potential. The establishment of naturally regenerating beech requires favourable conditions (Rodríguez-Ramírez et al., 2013) at an appropriate time and spatial pattern (Wagner et al., 2010, Barna, 2011). However, beech seedlings usually become established under a wide range of canopy openings (Sagheb-Talebi and Schütz, 2002). The survival and growth of regeneration is highly dependent on site and stand conditions, mainly due to the influence of stand structure on light availability (e.g. Madsen, 1994, Barna, 2008, Wagner et al., 2009), although saplings survive for long periods with low light exposure (Emborg, 1998, Modry et al., 2004).
The survival rate of different tree species during regeneration may differ significantly (Petritan et al., 2007) hence tree species composition may change noticeably during the regeneration period. The balance amongst species also depends on overstory and understory vegetation competition, browsing, the effect of pathogens, interactions with microfauna and microflora, and other abiotic factors such as late frost (Connell, 1990, Dovčiak and Brown, 2014).
Beech seedlings generally grow slower than most associated species (Beaudet and Messier, 1998), but survive longer during low or intermediate light availability (Logan, 1973, Nyland et al., 2006). Thus, forest management can change the composition of regeneration by regulating the degree of canopy density (Abe et al., 1995). However, beech may become dominant when reaching the young forest stage because of its high competitive power on beech-favourable sites. Height growth of beech increases after overstory cutting, even after growing under closed canopy conditions for long periods (Collet et al., 2001, Nagel et al., 2006, Emborg, 2007). Usually, stands with a dense understory of beech will lack other regeneration and the removal of beech is required to ensure the establishment of other species (Bohn and Nyland, 2003). Also, partial cutting of single trees or small patches may result in beech becoming more dominant (Collet et al., 2008).
Since beech has a high competitive ability at middle-elevation sites on mesotrophic soils, which may potentially exclude other species to survive, the aims of this study were to test the following questions:
- (1)
Does canopy openness control regeneration diversity or does beech regeneration remain dominant regardless of canopy openness?
- (2)
Can different intensities of strip shelterwood cutting influence the long term diversity of natural regeneration in submountain beech-dominated stands on mesotrophic soils?
To answer these questions, long-term data from plots established in 1989 was used to simulate varying intensities of canopy openness including clear cut.
Section snippets
Study area
The study was conducted between 2002 and 2012 in a submountain beech forest in the Kremnické vrchy Mts., western Carpathians, central Europe (48°38’N, 19°04’E, Fig. S1). The site is located on an exposed west facing slope with an inclination of 10–20° and the altitudinal range is between 470 and 510 m a.s.l. The site belongs to the moderately warm region and moderately warm and humid hilly land sub-region (Schieber, 2014). The mean annual air temperature and mean annual precipitation totals are
Changes in abundance and height growth of regeneration
Different shelterwood cutting and time since the initial cut had a significant effect on the number of regenerating individuals (Table 2). The highest density of regeneration occurred in the stand treated by a medium cut during the whole research (Fig. 1A) with an average of 6–10 individuals per m2. Year 2002 was an exception for C and L plots due to strong fructification of beech in the previous year. Regeneration density did not vary significantly in these stands during the research period,
Development of regeneration abundance
The abundance of natural regeneration is influenced mainly by the supply of seeds from the seed bank, sprouting success, survival and growth (Sagnard et al., 2007, Wagner et al., 2010). A large part of yearling seedlings in natural regeneration decreased with the decline of the number of fructifying trees in the parent stand (Barna, 2008). The most abundant fructifying trees were in the densest stands (C and L). High fructification of beech in these stands in 2001 triggered the rich
Conclusions
Results show that by applying various cutting methods, some attributes of subsequent stand replacement, emerging from natural regeneration, can change in the long-term, (in terms of successional evolution and long lasting change), but the interventions intensity only had a short-term impact on some of the attributes and with the stand gradually converging towards the same values.
With regards to the number of seedlings and regeneration density, the best results were achieved when applying a
Acknowledgements
This work was supported by the Scientific Grant Agency VEGA of the Ministry of Education of the Slovak Republic and of Slovak Academy of Sciences under Contract No. 2/0039/14. The work was also supported by the research project under the Contract No. NAZV, No. QJ1320230. Michal Bosela was supported by the Slovak Research and Development Agency under the project APVV-0273-11.
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