Éva SALAMON ALBERT, Ágnes CSISZÁR, Dénes BARTHA

Site conditions and functional traits affect regeneration dynamics of European hornbeam (Carpinus betulus L.) in forest canopy gaps

European hornbeam is a deciduous tree in mixed forest ecosystems, making a valuable contribution to species richness at a community level. We investigated early-phase regeneration of saplings in canopy gaps under contrasting ecological conditions by analyzing relationships between site parameters, leaf nutrients, and light-regulated carbon and water exchange functions at ecosystem and forest levels. Using plant response variables, we also detected functional soft traits and trade-offs. Species abundance proved to be affected by the elevation and slope of the sites, air humidity, precipitation during the growing period, as well as light-dependent and light-saturated rate of the assimilation function. Relationship between leaf potassium and transpiration proved to be the most effective plant functional trait across the sites. Under seasonally fluctuating, warm–dry and cold–semidry climate conditions with limited light availability, nutrient–gas exchange trade-offs modulated sapling development. Specifically during autumn, we detected extended leaf phenology in gas exchange functions under light-amplified warm and humid climate conditions. Our results suggest that natural regeneration of European hornbeam can be improved by lower and higher irradiation adjusted by plant functional traits, depending on site-specific and climate-related ecological parameters. These conclusions can provide a scientific basis for decision-making and practical implementations in forest management.

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Abe S, Masaki T, Nakashizuka T (1995). Factors influencing sapling composition in canopy gaps of a temperate deciduous forest. Vegetatio 120: 21-31.
Boratyński A (1996). Carpinus betulus. In: Roloff A, Weisgerber H, Lang UM, Stimm B, Schütt P, editors. Enzyklopädie der Holzgewächse: Handbuch und Atlas der Dendrologie III/2. Weinheim, DE: Wiley-Vch Verlag, pp. 1-12 (in German).
Borhidi A, Kevey B, Lendvai G (2012). Plant communities of Hungary. Budapest, Hungary: Akadémiai Kiadó.
Brang P, Spathelf P, Larsen JB, Bauhus J, Boncina A, Chauvin C, Drössler L, García-Güemes C, Heiri C, Kerr G et al. (2014). Suitability of close-to-nature silviculture for adapting temperate European forests to climate change. Forestry 87: 492-503.
Brokaw N, Busing RT (2000). Niche versus chance and tree diversity in forest gaps. Trends Ecol Evol 15: 183-188.
Busing RT, White PS (1997). Species diversity and small-scale disturbance in an old-growth temperate forest: a consideration of gap-partitioning concepts. Oikos 78: 562-568.
Canham CD (1989). Different responses to gaps among shadetolerant tree species. Ecology 70: 548-550.
Canham CD, Denslow JS, Platt WJ, Runkle JR, Spies TA, White PS (1990). Light regimes beneath closed canopies and tree-fall gaps in temperate and tropical forests. Can J Forest Res 20: 620-631.
Collins BS, Pickett STA (1987). Influence of canopy opening on the environment and herb-layer in a northern hardwoods forest. Vegetatio 70: 3-10.
Dreiss LM, Volin JC (2013). Influence of leaf phenology and site nitrogen on invasive species establishment in temperate deciduous forest understories. Forest Ecol Manag 296: 1-8.
EEA (2006). European forest types. Categories and types for sustainable forest management reporting and policy. Technical Report 9/2006, p. 111. Luxembourg City, Luxembourg: Office for Official Publications of the European Communities, [accessed 2018 March 28]. http://www.foresteurope.org/ docs/other_meetings/2006/wfc/WFC_4_eea_technical_ report_92006.pdf
EEA (2016). European forest ecosystems - State and trends. Technical Report 5/2016, p. 123. Luxembourg City, Luxembourg: Office for Official Publications of the European Communities, [accessed 2018 March 27]. https:// www.eea.europa.eu/ publications/european-forest-ecosystems
Field CB, Ball JT, Berry JA (1991). Photosynthesis: principles and field techniques. In: Ehleringer J, Mooney HA, Rundel PW, Pearcy RW, editors. Plant Physiological Ecology - Field Methods and Instrumentation. London, UK: Chapman and Hall, pp. 209-253.
Gálhidy L, Mihók B, Hagyó A, Rajkai K, Standovár T (2006). Effects of gap size and associated changes in light and soil moisture on the understory vegetation of a Hungarian beech forest. Plant Ecol 183: 133-145.
Kelemen K, Mihók B, Gálhidy L, Standovár, T (2012). Dynamic response of herbaceous vegetation to gap opening in a central European beech stand. Silva Fenn 46: 53-65.
Kevey B (2006). Oak-hornbeam forests of the floodplains of the Dráva river in Baranya, SW Hungary (article in Hungarian with an abstract in English). Nat Somogy 10: 41-71.
Kevey B (2008). Forest associations of Hungary (book in Hungarian with a summary in English). In: Bartha D, editor. Tilia XIV. Sopron, Hungary: University of West Hungary.
Kevey B (2012). Beech woods in the Eastern Mecsek Mountains [Helleboro odori-Fagetum (A.O. Horvát 1958) Soó & Borhidi in Soó 1960] (article in Hungarian with an abstract in English). e-Acta Nat Pannon 3: 27-48.
Kikuzawa K (1995). Leaf phenology as an optimal strategy for carbon gain in plants. Can J Bot 73: 158-163.
Klein T, Vitasse Y, Hoch G (2016). Coordination between growth, phenology and carbon storage in three coexisting deciduous tree species in a temperate forest. Tree Physiol 36: 847-855.
Köcher P, Horna V, Leuschner C (2013). Stem water storage in five coexisting temperate broad-leaved tree species: significance, temporal dynamics and dependence on tree functional traits. Tree Physiol 33: 817-832.
Leuzinger S, Zotz G, Asshoff R, Körner C (2005). Responses of deciduous forest trees to severe drought in Central Europe. Tree Physiol 25: 641-650.
Malcom DC, Mason WL, Clarke GC (2001). The transformation of conifer forests in Britain - regeneration, gap size and silvicultural systems. Forest Ecol Manag 151: 7-23.
Matuszkiewicz W (2003). Mixed oak-hornbeam forests (Carpinus betulus, Quercus robur, Q. petraea, Tilia cordata). In: Bohn U, Gollub G, Hettwer Ch, Neuhäuslová Z, Raus Th, Schlüter H, Weber H, editors. Map of the Natural Vegetation of Europe. Münster-Hiltrup, Germany: Bundesamt für Naturschutz, pp. 257-267.
McCarthy JW (2001). Gap dynamics of forest trees: A review with particular attention to boreal forests. Environ Rev 9: 1-59.
Messier C (1996). Managing light and understory vegetation in boreal and temperate broadleaf-conifer forests. In: Comeau PG, Thomas KD, editors. Silviculture of Temperate and Boreal Broadleaf-conifer Mixtures. Victoria, Canada: Victoria British Columbia Ministry of Forests, pp. 59-81.
Modry M, Hubeny D, Rejsek K (2004). Differential response of naturally regenerated European shade tolerant tree species to soil type and light availability. Forest Ecol Manag 188: 185-195.
Muscolo A, Sidari M, Mercurio R (2007). Influence of gap size on organic matter decomposition, microbial biomass and nutrient cycle in Calabrian pine (Pinus laricio Poiret) stands. Forest Ecol Manag 242: 412-418.
Muscolo A, Bagnato S, Sidari M, Mercurio R (2014). A review of the roles of forest canopy gaps. J Forestry Res 25: 725-736. Nock CA, Vogt RJ, Beisner BE (2016). Functional Traits. eLS 1-8.
Origin [CD-ROM] (2002). Release 7.0. Northampton, MA, USA: OriginLab Corporation. Accompanied by 1 user manual.
Paridari IC, Jalali SG, Sonboli A, Zarafshar M, Bruschi P (2013). Leaf macro- and micro-morphological altitudinal variability of Carpinus betulus in the Hyrcanian forest (Iran). J Forestry Res 24: 301-307.
Poulson TL, Platt WJ (1989). Gap light regimes influence canopy tree diversity. Ecology 70: 553-555.
Prescott CE, Hope GD, Blevins LL (2003). Effect of gap size on litter decomposition and soil nitrate concentrations in a high elevation spruce-fir forest. Can J Forest Res 33: 2210-2220.
Ritter E, Dalsgaard L, Einhorn KS (2005). Light, temperature and soil moisture regimes following gap formation in a semi-natural beech-dominated forest in Denmark. Forest Ecol Manag 206: 15-33.
Ritter E, Vesterdal L (2006). Gap formation in Danish beech (Fagus sylvatica) forests of low management intensity: soil moisture and nitrate in soil solution. Eur J Forest Res 125: 139-150.
Scharenbroch BC, Bockheim JG (2007). Impacts of forest gaps on soil properties and processes in old growth northern hardwoodhemlock forests. Plant and Soil 294: 219-233.
Scharenbroch BC, Bockheim JG (2008). Gaps and soil C dynamics in old growth northern hardwood-hemlock forests. Ecosystems 11: 426-441.
Schmidt W, Weitermeier M, Holzapfel C (1996). Vegetation dynamics in canopy gaps of a beech forest on limestone. The influence of the light gradient on species richness. Verh Ges Ökol 253-258.
Schmidt LM (2013). Extended leaf phenology in deciduous forest invaders: mechanisms of impact on native communities. J Veg Sci 24: 979-987.
Sikkema R, Caudullo G, de Rigo D (2016). Carpinus betulus in Europe: distribution, habitat, usage and threats. In: SanMiguel-Ayanz J, de Rigo D, Caudullo G, Houston Durrant T, Mauri A, editors. European Atlas of Forest Tree Species. Luxembourg City, Luxemburg: Publication Office of the European Union, pp. 74-75.
Statistica [download version] (2015). Data analysis software system. Release 12.5; Tulsa (OK): Dell Inc. Accompanied by 1 electronic user manual.
Stojnić S, Pekeć S, Kebert M, Pilipović A, Stojanović D, Stojanović M, Orlović S (2016). Drought effects on physiology and biochemistry of pedunculate oak (Quercus robur L.) and hornbeam (Carpinus betulus L.) saplings grown in urban area of Novi Sad, Serbia. SEEFOR 7: 57-63.
Vajari AK, Jalilvand H, Pourmajidian MR, Espahbodi K, Moshki A (2012). Effect of canopy gap size and ecological factors on species diversity and beech seedlings in managed beech stands in Hyrcanian forests. J Forestry Res 23: 217-222.
Violle C, Navas ML, Vile D, Kazakou E, Fortunel C, Hummel I, Garnir E (2007). Let the concept of trait be functional! Oikos 116: 882-892.
Vitasse YS, Delzon E, Dufrene J, Pontailler J, Louvet A, Kremer A, Michalet R (2009). Leaf phenology sensitivity to temperature in European trees: Do within-species populations exhibit similar responses? Agr Forest Meteorol 149: 735-744.
Zirlewagen D, von Wilpert K (2001). Modeling water and ion fluxes in a highly structured, mixed-species stand. Forest Ecol Manag 143: 27-37.