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iForest - Biogeosciences and Forestry

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The maternal environment of European beech (Fagus sylvatica L.) affects intrapopulation variability in seed traits and germination

Vladan Popović (1)   , Aleksandar Lučić (1), Mladen Ivanković (2), Branislav Cvjetković (3), Gregor Božič (4), Vlatko Andonovski (5), Danijela Miljković (6)

iForest - Biogeosciences and Forestry, Volume 19, Issue 2, Pages 114-121 (2026)
doi: https://doi.org/10.3832/ifor4806-018
Published: Apr 10, 2026 - Copyright © 2026 SISEF

Research Articles


Interpopulation variation was investigated using seed samples originating from twenty-six European beech (Fagus sylvatica L.) populations across the Balkan Peninsula, a part of the species’ distribution range characterized by high ecological heterogeneity in key climatic factors, such as temperature (5.8-10.6 °C), precipitation (648-1632 mm), and elevation (185-1410 m a.s.l.). The statistical significance of intrapopulation differences was confirmed by analysis of variance (ANOVA) for all seed traits analyzed: seed weight (g), length (mm), width (mm), thickness (mm), eccentricity and flatness indices, and germination capacity (%). Multivariate principal component analysis (PCA) was applied to examine seed traits in relation to environmental variables of the maternal site, such as mean temperature and precipitation in September and October (the seed maturation period), revealing distinct patterns of relationships among the variables studied. Seed traits were significantly positively correlated with mean temperatures of the maternal site in September and October, indicating that temperature during the seed-filling period affects seed mass. Germination capacity was associated with precipitation during the same period, though the correlation coefficient was not statistically significant; a shorter vector length in the PC biplot suggests a weaker contribution to population separation. Elevation of the site of origin showed a significant negative correlation with temperature, precipitation, and seed traits. Agglomerative hierarchical clustering analysis identified three distinct population clusters. Higher temperature and precipitation values did not necessarily result in higher seed trait values or higher germination percentages. The population with the highest seed mass exhibited the lowest germination capacity (32%) during seed maturation under the lowest precipitation. Conversely, the population characterized by the lowest seed mass showed a higher germination rate of 68% in environments with high precipitation. These results provide valuable insights into the reproductive ecology of European beech, suggesting that other factors beyond those analyzed here may have a more substantial influence on seed germination. The variation in seed traits across habitats that are either drier and hotter or colder and wetter, along the elevation gradient of the studied populations, paves the way for future research and breeding efforts to enhance the species’ survival and reproductive success amid anticipated climate change scenarios.

  Keywords


Seed Traits, Seed Germination, European Beech, Fagus sylvatica L., Environmental and Genetic Variation, Southeast Europe

Authors’ address

(1)
Vladan Popović 0000-0003-4326-3364
Aleksandar Lučić 0000-0002-4473-1791
Department of Genetics, Plant Breeding, Seed and Nursery Production; Institute of Forestry, Kneza Višeslava 3, 11030 Belgrade (Serbia)
(2)
Mladen Ivanković 0000-0003-1198-9902
Department for Genetics, Forest Tree Breeding and Seed Science, Croatian Forest Research Institute, Cvjetno naselje 41, 10450 Jastrebarsko (Croatia)
(3)
Branislav Cvjetković 0000-0002-3589-5516
Faculty of Forestry, University of Banja Luka, Bulevar vojvode Petra Bojovića 1A, 78000 Banja Luka, Republic of Srpska (Bosnia and Herzegovina)
(4)
Gregor Božič 0000-0002-5595-2979
Department of Forest Physiology and Genetics, Slovenian Forestry Institute, Večna pot 2, 1000 Ljubljana (Slovenia)
(5)
Vlatko Andonovski 0000-0001-9697-5854
University Ss. Cyril and Methodius, Hans Em Faculty of Forest Sciences, Landscape Architecture and Environmental Engineering, 16 Makedonska brigada 1, 1000 Skopje (North Macedonia)
(6)
Danijela Miljković 0000-0003-1781-6658
Department of Evolutionary Biology, University of Belgrade, Institute for Biological Research “Siniša Stanković”, National Institute of the Republic of Serbia, Bulevar despota Stefana 142, 11060 Belgrade (Serbia)

Corresponding author

 
Vladan Popović
vladanpop79@gmail.com

Citation

Popović V, Lučić A, Ivanković M, Cvjetković B, Božič G, Andonovski V, Miljković D (2026). The maternal environment of European beech (Fagus sylvatica L.) affects intrapopulation variability in seed traits and germination. iForest 19: 114-121. - doi: 10.3832/ifor4806-018

Academic Editor

Rafael Da Silveira Bueno

Paper history

Received: Jan 26, 2025
Accepted: Apr 09, 2026

First online: Apr 10, 2026
Publication Date: Apr 30, 2026
Publication Time: 0.03 months

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(1)
Allen CD, Breshears DD, McDowell NG (2015)
On the underestimation of global vulnerability to tree mortality and forest die-off from hotter drought in the Anthropocene. Ecosphere 6: 1-55.
CrossRef | Gscholar
(2)
Bajocco S, Ferrara C, Bascietto M, Alivernini A, Chirichella R, Cutini A, Chianucci F (2021)
Characterizing the climatic niche of mast seeding in beech: evidence of trade-offs between vegetation growth and seed production. Ecological Indicators 121: 107139.
CrossRef | Gscholar
(3)
Baskin CC, Baskin JM (2014)
Seeds: Ecology, biogeography, and evolution of dormancy and germination (2nd edn). Academic Press/Elsevier, San Diego, CA, USA. pp. 277-373.
Gscholar
(4)
Bezdečková L, Matejka K (2015)
The quality and the depth of dormancy of beechnuts in individual stand groups with varying climatic conditions within a single unit of approval. Journal of Forest Science 61 (9): 382-392.
CrossRef | Gscholar
(5)
Bu H, Chen X, Xu X, Liu K, Jia P, Du G (2008)
Seed mass and germination in an alpine meadow on the eastern Tsinghai-Tibet Plateau. Plant Ecology 197 (1): 69-76.
CrossRef | Gscholar
(6)
Cerabolini BRM, Ceriani M, Caccianiga R, Andreis D, Raimondi B (2003)
Seed size, shape and persistence in soil: a test on Italian flora from Alps to Mediterranean coasts. Seed Science Research 13 (1): 75-85.
CrossRef | Gscholar
(7)
Cochrane A, Hoyle G, Yates C, Wood J, Nicotra A (2015)
Will among-population variation in seed traits improve the chance of species persistence under climate change? Global Ecology and Biogeography 24 (11): 1211-1224.
CrossRef | Gscholar
(8)
Czúcz B, Galhidy L, Matyas C (2011)
Present and forecasted xeric climatic limits of beech and sessile oak distribution at low altitudes in Central Europe. Annals of Forest Science 68 (1): 99-108.
CrossRef | Gscholar
(9)
Despland E, Houle G (1997)
Climate influences on growth and reproduction of Pinus banksiana (Pinaceae) at the limit of the species distribution in eastern North America. American Journal of Botany 84 (7): 928-937.
CrossRef | Gscholar
(10)
Drvodelić D, Oršanić M, Paulić V, Roman M (2011)
Morphological-biological properties of fruit and seed of beech (Fagus sylvatica L.) growing at different altitudes. Glas Šum Pokuse 44: 1-19.
Gscholar
(11)
Falk W, Hempelmann N (2013)
Species favourability shift in Europe due to climate change: a case study for Fagus sylvatica L. and Picea abies (L.) Karst. based on an ensemble of climate models. Journal of Climatology 213: 787250.
CrossRef | Gscholar
(12)
Ferus P, Barta M, Konôpková J, Turčeková S, Manka P, Biben T (2013)
Diversity in honey locust (Gleditsia triacanthos L.) seed traits across Danube basin. Folia Oecologica 40 (2): 163-169.
Online | Gscholar
(13)
Friedman J, Middleton TE, Rubin MJ (2019)
Environmental heterogeneity generates intrapopulation variation in life-history traits in an annual plant. New Phytologist 224 (3): 1171-1183.
CrossRef | Gscholar
(14)
Gavranović A, Bogdan S, Lanšćak M, Cehulić I, Ivanković M (2018)
Seed yield and morphological variations of beech nuts in four European beech (Fagus sylvatica L.) populations in Croatia. South-East European Forestry 9 (1): 17-27.
CrossRef | Gscholar
(15)
Gavranović Markić A, Vujnović Z, Kičić M, Ivanković M (2024)
Seed quantity and quality variation in European beech (Fagus sylvatica L.): a comparative analysis of different crop years. South-East European Forestry 15 (1): 1-12.
CrossRef | Gscholar
(16)
Gradečki M, Postenjak K, Littvay T (2003)
Neke morfološke i fiziološke značajke sjemena bukve [Some morphological and physiological characteristics of beech seed]. In: Proceedings of “8 Hrvatski Biološki Kongres” (Besendorfer V, Kopjar N eds). Hrvatsko Biološko Društvo 1885, Zagreb, Hrvatska, pp. 196-197. [in Croatian]
Gscholar
(17)
Griesbauer H, DeLong SC, Rogers B, Foord V (2021)
Growth sensitivity to climate varies with soil moisture regime in spruce-fir forests in central British Columbia. Trees 35: 649-669.
CrossRef | Gscholar
(18)
Hamann A, Wang T, Spittlehouse DL, Murdock TQ (2013)
A comprehensive, high-resolution database of historical and projected climate surfaces for western North America. Bulletin of the American Meteorological Society 94 (9): 1307-1309.
CrossRef | Gscholar
(19)
Hilton GM, Packham JR (2003)
Variation in the masting of common beech (Fagus sylvatica L.) in northern Europe over two centuries (1800-2001). Forestry 76 (3): 319-328.
CrossRef | Gscholar
(20)
ISTA (1996)
International rules for seed testing. Seed Science and Technology 13: 299-513.
Gscholar
(21)
Jump AS, Mátyás C, Peñuelas J (2009)
The altitude-for-latitude disparity in the range retractions of woody species. Trends in Ecology and Evolution 24 (12): 694-701.
CrossRef | Gscholar
(22)
Lebourgeois F, Delpierre N, Dufrêne E, Cecchini S, Macé S, Croisé L, Nicolas M (2018)
Assessing the roles of temperature, carbon inputs and airborne pollen as drivers of fructification in European temperate deciduous forests. European Journal of Forest Research 137 (3): 349-365.
CrossRef | Gscholar
(23)
Leimu R, Mutikainen P, Koricheva J, Fischer M (2006)
How general are positive relationships between plant population size, fitness and genetic variation? Journal of Ecology 94 (5): 942-952.
CrossRef | Gscholar
(24)
Madrigal-González J, Ballesteros-Cánovas JA, Herrero A (2017)
Forest productivity in southwestern Europe is controlled by coupled North Atlantic and Atlantic Multidecadal Oscillations. Nature Commununications 8 (1): 2222.
CrossRef | Gscholar
(25)
Matías L, Jump AS (2014)
Impacts of predicted climate change on recruitment at the geographical limits of Scots pine. Journal of Experimental Botany 65 (1): 299-310.
CrossRef | Gscholar
(26)
Mazza G, Monteverdi MC, Altieri S, Battipaglia G (2024)
Climate-driven growth dynamics and trend reversal of Fagus sylvatica L. and Quercus cerris L. in a low-elevation beech forest in Central Italy. Science of The Total Environment 908: 168250.
CrossRef | Gscholar
(27)
Mishra Y, Rawat R, Rana PK, Sonkar MK, Mohammad N (2014)
Effect of seed mass on emergence and seedling development in Pterocarpus marsupium Roxb. Journal of Forestry Research 25 (2): 415-418.
CrossRef | Gscholar
(28)
Muffler L, Schmeddes J, Weigel R (2021)
High plasticity in germination and establishment success in the dominant forest tree Fagus sylvatica across Europe. Global Ecology and Biogeography 30 (8): 1583-1596.
CrossRef | Gscholar
(29)
Müller-Haubold H, Hertel D, Leuschner C (2015)
Climatic drivers of mast fruiting in European beech and resulting C and N allocation shifts. Ecosystems 18 (6): 1083-1100.
CrossRef | Gscholar
(30)
Murray BR, Brown AHD, Dickman CR, Crowther MS (2004)
Geographical gradients in seed mass in relation to climate. Journal of Biogeography 31 (3): 379-388.
CrossRef | Gscholar
(31)
Naudiyal N, Wang J, Ning W, Gaire NP, Peili S, Yanqiang W, Ning S (2021)
Potential distribution of Abies, Picea, and Juniperus species in the sub-alpine forest of Minjiang headwater region under current and future climate scenarios and its implications on ecosystem services supply. Ecological Indicators 121 (1): 107131.
CrossRef | Gscholar
(32)
Pakeman RJ, Eastwood A (2013)
Shifts in functional traits and species’ niche space response to climate change in the UK. Journal of Vegetation Science 24 (5): 865-876.
CrossRef | Gscholar
(33)
Pérez-Ramos IM, Marañón T (2009)
Effects of waterlogging on seed germination of three Mediterranean oak species: ecological implications. Acta Oecologica 35 (3): 422-428.
CrossRef | Gscholar
(34)
Piovesan G, Biondi F, Di Filippo A, Alessandrini A, Maugeri M (2008)
Drought-driven growth reduction in old beech (Fagus sylvatica) forests of the central Apennines, Italy. Global Change Biology 14 (6): 1265-1281.
CrossRef | Gscholar
(35)
Popović V, Sijačić-Nikolić M, Ristić D (2015)
Analiza kvaliteta i morfometrijskuh karakteristika semena bukve (Fagus moesiaca/Domin, Maly/Czeczott) u Srbiji [Variability of morphometric characteristics of seed and height of one-year-old seedlings of different populations of beech (Fagus moesiaca/Domin, Maly/Czeczott) in Serbia]. Šumarstvo 1-2: 109-120. [in Bosnian]
Gscholar
(36)
Pšidová E, Ditmarová L, Jamnická G, Kurjak D, Majerová J, Cazajkowski T, Bolte A (2015)
Photosynthetic response of beech seedlings of different origins to water deficit. Photosynthetica 53 (2): 187-194.
CrossRef | Gscholar
(37)
Sánchez-Velásquez LR, Del Rosario Pineda-López M, Ibarra-Zavaleta SP, López-Serrano Y (2021)
Fir forest demography using matrix projections: anomaly precipitation due to climatic change decreases population viability. Forest Ecology and Management 482: 118845.
CrossRef | Gscholar
(38)
SAS Institute Inc. (2011)
The SAS System for Windows, release 9. 3. SAS Institute, Cary, NC, USA.
Gscholar
(39)
Shu K, Qi Y, Chen F, Meng Y, Luo X, Shuai H, Zhou W, Ding J, Du J, Liu J, Yang F, Wang Q, Liu W, Yong T, Wang X, Feng Y, Yang W (2017)
Salt stress represses soybean seed germination by negatively regulating GA biosynthesis while positively mediating ABA biosynthesis. Frontiers in Plant Science 8: 938.
CrossRef | Gscholar
(40)
Smaill SJ, Clinton PW, Allen RB, Davis MR (2011)
Climate cues and resources interact to determine seed production by a masting species. Journal of Ecology 99 (3): 870-877.
CrossRef | Gscholar
(41)
Smelkova L (2002)
Coating of seeds of forest woody species and its impact on the germination and growth of seedlings. Šumarski List 125 (5/6): 243-248.
Online | Gscholar
(42)
Stojnić S, Suchocka M, Benito-Garzón M, Torres-Ruiz JM, Cochard H, Bolte A, Cocozza C, Cvjetković B, de Luis M, Martinez-Vilalta J, Ræbild A, Tognetti R, Delzon S (2018)
Variation in xylem vulnerability to embolism in European beech from geographically marginal populations. Tree Physiology 38 (2): 173-185.
CrossRef | Gscholar
(43)
Vacchiano G, Hackett-Pain A, Turco M, Motta R, Maringer J, Conedera M, Drobyshev I, Ascoli D (2017)
Spatial patterns and broad-scale weather cues of beech mast seeding in Europe. New Phytologist 215 (2): 595-608.
CrossRef | Gscholar
(44)
Varsamis G, Merou T, Takos I, Malesios C, Manolis A, Papageorgiou A (2020)
Seed adaptive traits of Fagus sylvatica populations in Northeastern Greece. Forest Science 66 (4): 403-415.
CrossRef | Gscholar
(45)
Von Wühlisch G (2008)
European beech EUFORGEN Technical Guidelines for Genetic Conservation and Use 6, Bioversity International, Rome, Italy, pp. 6.
Online | Gscholar
(46)
Wagner JE (2011)
Forestry economics: a managerial approach (Vol. 3). Routledge, Abingdon, Oxon, UK and New York, NY, USA, pp. 382.
Gscholar
(47)
Yilmaz M (2010)
Is there a future for the isolated oriental beech (Fagus orientalis Lipsky) forests in southern Turkey? Acta Silvatica et Lignaria Hungarica 6: 111-114.
Gscholar
 

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