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Effects of light transmittance on growth and biomass of understory seedlings in mixed pine-beech forests

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Abstract

Seedling growth as well as aboveground and belowground biomass allocation is mostly influenced by Light Transmittance (LT) (%) through the canopy. The knowledge of how understory light conditions affect seedling growth and biomass of different species in mixed forests is not well documented. Thus, it is essential to quantify the effects of light on the growth and biomass of understory seedlings. Given their advantages over pure forests, these quantitative understandings are especially crucial in mixed forests with species whose light demand and shade tolerance vary. This research examined the growth responses of natural-origin Scots pine (Pinus sylvestris L.), black pine (Pinus nigra Arnold.) and Oriental beech (Fagus orientalis L.) seedlings to LT (%) through the canopy in their mixed stands. Linear mixed-effect models were utilized to examine the influence of LT (%) on the seedlings. Moreover, allometric equations for estimating the aboveground biomass and belowground biomass of seedlings were developed for each species. Seedling height, and aboveground and belowground biomass after five years of germination were most significantly affected by LT (%) and tree species in the mixed pine-beech stand (p < 0.001). Biomass models for each species included different variable combinations of seedling height, root-collar diameter, LT (%), and their interactions. This study demonstrates the importance of canopy structure and overstory disturbances for the maintenance of mixed pine-beech forests since canopy structure significantly contributes to the understory light environment.

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Data availability

The datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request.

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References

  • Addo-Danso SD, Prescott CE, Smith AR (2016) Methods for estimating root biomass and production in forest and woodland ecosystem carbon studies: a review. For Ecol Manag 359:332–351

    Article  Google Scholar 

  • Akbenar KT, Keshavarz M (2005) Influence of light condition on quality and quantity characteristics of beech (Fagus orientalis Lipsky) sapling in north of Iran. Asian J Plant Sci 4:161–163

    Google Scholar 

  • Annighöfer P, Ameztegui A, Ammer C, Balandier P, Bartsch N, Bolte A, Mund M (2016) Species-specific and generic biomass equations for seedlings and saplings of European tree species. Eur J for Res 135(2):313–329

    Article  Google Scholar 

  • Annighöfer P, Mund M, Seide D, Ammer C, Ameztegui A, Balandier P, Bebre I, Coll L, Collet C, Hamm T, Huth F (2022) Examination of aboveground attributes to predict belowground biomass of young trees. For Ecol Manag 505:119942

    Article  Google Scholar 

  • Arnold TW (2010) Uninformative parameters and model selection using Akaike’s information criterion. J Wildl Manag 74(6):1175–1178

    Article  Google Scholar 

  • Assenac G (2000) Interactions between forest stands and microclimate: ecophysiological aspects and consequences for silviculture. Ann for Sci 57:287–301

    Article  Google Scholar 

  • Atik A (2013) Effects of planting density and treatment with vermicompost on the morphological characteristics of oriental beech (Fagus orientalis Lipsky.). Compost Sci Util 21:87–98

    Article  Google Scholar 

  • Babaei F, Jalali SG, Sohrabi H, Shirvany A (2017) Variability in leaf and crown morphology correlated with light availability in five natural populations of Quercus castaneifolia CA Mey. J for Sci 63(6):275–281

    Article  Google Scholar 

  • Bachofen C, Wohlgemuth T, Moser B (2019) Biomass partitioning in a future dry and CO2 enriched climate: shading aggravates drought effects in Scots pine but not European black pine seedlings. J Appl Ecol 56(4):866–879

    Article  CAS  Google Scholar 

  • Bayala J, Dianda M, Wilson J, Ouedraogo S, Sanon K (2009) Predicting field performance of five irrigated tree species using seedling quality assessment in Burkina Faso. West Afr New for 38(3):309–322

    Google Scholar 

  • Bonser SP, Aarssen LW (1994) Plastic allometry in young sugar maple (Acer saccharum): adaptive responses to light availability. Am J Bot 81:400–406

    Article  Google Scholar 

  • Çalışkan A (1992) Karabük-Büyükdüz Araştırma ormanında sarıçam (Pinus sylvestris L.)-Göknar (Abies bornmülleriana Mattf.)-Kayın (Fagus orientalis Lipsky) karışık meşcerelerinde büyüme ilişkileri ve gerekli silvikültürel işlemler. J Fac for Istanb Univ 42(2):184–209

    Google Scholar 

  • Castro J, Zamora R, Hódar JA, Gómez JM (2004) Seedling establishment of a boreal tree species (Pinus sylvestris) at its southernmost distribution limit: consequences of being in a marginal Mediterranean habitat. J Ecol 92(2):266–277

    Article  Google Scholar 

  • Čater M, Schmid I, Kazda M (2013) Instantaneous and potential radiation effect on underplanted European beech below Norway spruce canopy. Eur J for Res 132(1):23–32

    Article  Google Scholar 

  • Chiarucci AJBW, Wilson JB, Anderson BJ, De Dominicis V (1999) Cover versus biomass as an estimate of species abundance: Does it make a difference to the conclusions? J Veg Sci 10(1):35–42

    Article  Google Scholar 

  • Collet C, Lanter O, Pardos M (2001) Effects of canopy opening on height and diameter growth in naturally regenerated beech seedlings. Ann for Sci 58(2):127–134

    Article  Google Scholar 

  • Danescu A, Albrecht AT, Bauhus J (2016) Structural diversity promotes productivity of mixed, uneven-aged forests in southwestern Germany. Oecologia 182:319–333

    Article  PubMed  Google Scholar 

  • Daryaei A, Sohrabi H, Puerta-Piñero C (2019) How does light availability affect the aboveground biomass allocation and leaf morphology of saplings in temperate mixed deciduous forests? New for 50(3):409–422

    Article  Google Scholar 

  • DeYoung J (2016) Forest measurements: an applied approach. Open oregon educational resources, Oregon- USA

  • Domenicano S, Coll L, Messier C, Berninger F (2011) Nitrogen forms affect root structure and water uptake in the hybrid poplar. New for 42:347–362

    Article  Google Scholar 

  • Dormann CF, Elith J, Bacher S, Buchmann C, Carl G, Carré G et al. (2013) Collinearity: a review of methods to deal with it and a simulation study evaluating their performance. Ecography 36(1):27–46

    Article  Google Scholar 

  • Gaudio N, Balandier P, Perret S, Ginisty C (2011) Growth of understorey Scots pine (Pinus sylvestris L.) saplings in response to light in mixed temperate forest. Forestry 84(2):187–195

    Article  Google Scholar 

  • Genç M (2020) Orman bakımı: Asli Orman Ağacı Türlerimizin Saf ve Karışık Meşcerelerinin Bakımı. Giresun, Musa Genc Kitaplığı, p 216 (in Turkish)

    Google Scholar 

  • Gerelbaatar S, Baatarbileg N, Batsaikhan G (2015) Density and growth performance of Scots pine (Pinus sylvestris L.) understory saplings in logged forests of Tujiin Nars national park in the western Khentii mountains, case study from northern Mongolia. Glob Adv Res J Agric Sci 4(1):17–25

    Google Scholar 

  • Grubb PJ, Lee WG, Kollmann J, Wilson JB (1996) Interaction of irradiance and soil nutrient supply on growth of seedlings of ten European tall-shrub species and Fagus sylvatica. J Ecol 84:827–840

    Article  Google Scholar 

  • Hale SH (2003) The effect of thinning intensity on the below-canopy light environment in a Sitka spruce plantation. For Ecol Manag 179:341–349

    Article  Google Scholar 

  • Hallikainen V, Hyppönen M, Hyvönen J, Niemelä J (2007) Establishment and height development of harvested and naturally regenerated Scots pine near the timberline in North-East Finnish Lapland. Silva Fenn 41(1):71–88

    Article  Google Scholar 

  • Huang Y, Chen Y, Castro-Izaguirre N, Baruffol M, Brezzi M, Lang A, Schmid B (2018) Impacts of species richness on productivity in a large-scale subtropical forest experiment. Science 362:80–83

    Article  PubMed  CAS  Google Scholar 

  • Humbert L, Gagnon D, Kneeshaw D, Messier C (2007) A shade tolerance index for common understory species of northeastern North America. Ecol Indic 7:195–207

    Article  Google Scholar 

  • Hyppönen M, Hallikainen V, Niemelä J, Rautio P (2013) The contradictory role of understory vegetation on the success of Scots pine regeneration. Silva Fenn 47(1):903

    Article  Google Scholar 

  • Juntunen V, Neuvonen S (2006) Natural regeneration of Scots pine and Norway spruce close to the timberline in Northern Finland. Silva Fenn 40(3):443–458

    Article  Google Scholar 

  • Kara F, Topaçoğlu O (2018a) Effects of canopy structure on growth and aboground/belowground biomass of seedlings in uneven-aged Trojan fir stands. Cerne 24(4):312–322

    Article  Google Scholar 

  • Kara F, Topaçoğlu O (2018b) Influence of stand density and canopy structure on the germination and growth of Scots pine (Pinus sylvestris L.) seedlings. Environ Monit Assess 190(12):1–10

    Article  CAS  Google Scholar 

  • Kara F, Loewenstein EF, Lhotka JM, Kush JS (2018) A Gingrich style stocking chart for longleaf pine (Pinus palustris Mill.) forests. For Sci 64(3):307–315

    Google Scholar 

  • Köseoğlu E, Kara F (2019) Comparison of the influence of some stand structural parameters on black pine seeling density and growth. Turk J for 20(4):305–311

    Article  Google Scholar 

  • Kuusipalo J (1983) Mustikan varvuston biomassamäärän vaihtelusta erilaisissa metsiköissä. Silva Fenn 17(3):245–257 (in Finnish)

    Article  Google Scholar 

  • Lhotka JM, Loewenstein EF (2008) Influence of canopy structure on the survival and growth of underplanted seedlings. New for 35(1):89–104

    Article  Google Scholar 

  • Macaroğlu K (2011) Study on biomass carbon sequestration capacities of mixed stands in Bartın. Master thesis. Bartın University, Bartın, p 116

    Google Scholar 

  • Madsen P (1994) Growth and survival of Fagus sylvatica seedlings in relation to light intensity and soil water content. Scand J for Res 9:316–322

    Article  Google Scholar 

  • Mason WL, Edwards C, Hale SE (2004) Survival and early seedling growth of conifers with different shade tolerance in a Sitka spruce spacing trial and relationship to understory light climate. Silva Fenn 38(4):357–370

    Article  Google Scholar 

  • Mediavilla S, Escudero A (2010) Differences in biomass allocation patterns between saplings of two co-occurring Mediterranean oaks as reflecting different strategies in the use of light and water. Eur J for Res 129:697–706

    Article  Google Scholar 

  • Mercurio R, Mallamaci C, Muscolo A, Sidari M (2009) Effetti della dimensione delle buche sulla rinnovazione naturale in rimboschimenti di pino nero. For J Silvic for Ecol 6(5):312

    Google Scholar 

  • Messier C, Nikinmaa E (2000) Effects of light availability and sapling size on the growth, biomass allocation, and crown morphology of understory sugar maple, yellow birch, and beech. Ecoscience 7(3):345–356

    Article  Google Scholar 

  • Messier C, Parent S, Bergeron Y (1998) Effects of overstory and understory vegetation on the understory light environment in mixed boreal forests. J Veg Sci 9(4):511–520

    Article  Google Scholar 

  • Messier C, Doucet R, Ruel JC, Claveau Y, Kelly C, Lechowicz MJ (1999) Functional ecology of advance regeneration in relation to light in boreal forests. Can J for Res 29:812–823

    Article  Google Scholar 

  • Modrzyński J, Chmura DJ, Tjoelker MG (2015) Seedling growth and biomass allocation in relation to leaf habit and shade tolerance among 10 temperate tree species. Tree Physiol 35(8):879–893

    Article  PubMed  Google Scholar 

  • Muukkonen P, Mäkipää R, Laiho R, Minkkinen K, Vasander H, Finér L (2006) Relationship between biomass and percentage cover in understorey vegetation of boreal coniferous forests. Silva Fenn 40:231–245

    Article  Google Scholar 

  • Niinemets Ü (1996) Plant growth-form alters the relationship between foliar morphology and species shade-tolerance ranking in temperate woody taxa. Vegetatio 124:145–153

    Article  Google Scholar 

  • Odabaşı T, Calışkan A, Bozkus HF (2004) Silvikültür tekniği. İstanbul University Publications, İstanbul, p 314 (in Turkish)

    Google Scholar 

  • Pádua CBV, Chiaravalotti R (2012) Silviculture and biodiversity. Writings of the dialogue, vol 4. Apremavi, Rio do Sul, SC, pp. 68. ISBN 978-85-88733-09-1

  • Pardos M, Montes F, Aranda I, Cañellas I (2007) Influence of environmental conditions on germinant survival and diversity of Scots pine (Pinus sylvestris L.) in central Spain. Eur J for Res 126(1):37–47

    Article  Google Scholar 

  • Parhizkar P, Sagheb-Talebi K, Mataji A, Nyland R, Namiranian M (2011) Silvicultural characteristics of Oriental beech (Fagus orientalis Lipsky) regeneration under different RLI and positions within gaps. Forestry 84:177–185

    Article  Google Scholar 

  • Petriţan AM, Von Lüpke B, Petriţan IC (2009) Influence of light availability on growth, leaf morphology and plant architecture of beech (Fagus sylvatica L.), maple (Acer pseudoplatanus L.) and ash (Fraxinus excelsior L.) saplings. Eur J for Res 128:61–74

    Article  Google Scholar 

  • Petrovska R, Brang P, Gessler A, Bugmann H, Hobi ML (2021) Grow slowly, persist, dominate—explaining beech dominance in a primeval forest. Ecol Evol 11(15):10077–10089

    Article  PubMed  PubMed Central  Google Scholar 

  • Pretzsch H, del Río M, Ammer C, Avdagic A, Barbeito I, Bielak K, Bravo-Oviedo A (2015) Growth and yield of mixed versus pure stands of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) analysed along a productivity gradient through Europe. Eur J for Res 134(5):927–947

    Article  Google Scholar 

  • Puttonen P (1997) Looking for the ‘“silver bullet”’—can one test do it all? New for 13:9–27

    Article  Google Scholar 

  • R Development Core Team (2021) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria

  • Röttgermann M, Steinlein T, Beyschlag W, Dietz H (2000) Linear relationships between aboveground biomass and plant cover in low open herbaceous vegetation. J Veg Sci 11(1):145–148

    Article  Google Scholar 

  • Sevillano I, Short I, Grant J, O’Reilly C (2016) Effects of light availability on morphology, growth and biomass allocation of Fagus sylvatica and quercus robur seedlings. For Ecol Manag 374:11–19

    Article  Google Scholar 

  • Shahrokhzadeh U, Sohrabi H, Copenheaver CA (2015) Aboveground biomass and leaf area equations for three common tree species of Hyrcanian temperate forests in northern Iran. Botany 93:663–670

    Article  CAS  Google Scholar 

  • Snipes M, Taylor DC (2014) Model selection and Akaike information criteria: an example from wine ratings and prices. Wine Econ Policy 3(1):3–9

    Article  Google Scholar 

  • Tardós P, Lucas-Borja ME, Beltrán M, Onkelinx T, Piqué M (2019) Composite low thinning and slash burning treatment enhances initial Spanish black pine seedling recruitment. For Ecol Manag 433:1–12

    Article  Google Scholar 

  • Tíscar P, Linares J (2014) Large-scale regeneration patterns of Pinus nigra subsp. salzmannii: poor evidence of increasing facilitation across a drought gradient. Forests 5(1):1–20

    Article  Google Scholar 

  • Topaçoğlu O, Genç E (2019) Forest Edge Effects on Seedlings in Mixed Oriental Beech (Fagus orientalis Lipsky)-Scots Pine (Pinus sylvestris L.) Stands. Appl Ecol Environ Res 17(2):2219–2231

    Article  Google Scholar 

  • Wagner S, Collet C, Madsen P, Nakashizuka T, Nyland RD, Sagheb-Talebi K (2010) Beech regeneration research: from ecological to silvicultural aspects. For Ecol Manag 259:2172–2182

    Article  Google Scholar 

  • Walters MB, Reich PB (2000) Seed size, nitrogen supply, and growth rate affect tree seedling survival in deep shade. Ecology 81(7):1887–1901

    Article  Google Scholar 

  • Welander NT, Ottosson B (1998) The influence of shading on growth and morphology in seedlings of Quercus robur L. and Fagus sylvatica L. For Ecol Manag 107(1–3):117–126

    Article  Google Scholar 

  • Yamashita N, Okuda S, Suwa R, Lei TT, Tobita H, Utsugi H, Kajimoto T (2016) Impact of leaf removal on initial survival and growth of container-grown and bare-root seedlings of Hinoki cypress (Chamaecyparis obtusa). For Ecol Manag 370:76–82

    Article  Google Scholar 

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Acknowledgements

The authors would like to thank Kastamonu Regional Directorate of Forestry for providing access to the study area for this research. Moreover, the authors would acknowledge Emre Genç, Erol Kaya and Abdurrahman Göksu for their help during the fieldworks.

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This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors.

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FK conceived the idea, developed the framework, collected data and compiled the literature, and worked with the writing and editing. FK conceptualized the study, collected the data, performed the statistical analyses, created figures and wrote the whole manuscript.

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Correspondence to Ferhat Kara.

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Communicated by: Chirstian Ammer.

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Kara, F. Effects of light transmittance on growth and biomass of understory seedlings in mixed pine-beech forests. Eur J Forest Res 141, 1189–1200 (2022). https://doi.org/10.1007/s10342-022-01501-4

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