European Journal of Forest Research

, Volume 134, Issue 6, pp 1095–1108 | Cite as

Climate modifies tree interactions in terms of basal area growth and mortality in monospecific and mixed Fagus sylvatica and Pinus sylvestris forests

  • Sonia CondésEmail author
  • Miren del Río
Original Paper


Net interactions between trees vary depending on environmental conditions and other factors such as stand density, age, or between-species complementarity and/or facilitation. According to the stress gradient hypothesis, positive or facilitative interactions are more frequent in high-stress environments whereas negative or competitive interactions occur in benign environments, although recent studies highlight the influence of species composition, type of stress, ontogeny, etc. on the interaction–stress gradient pattern. The aim of this paper is to analyze whether site climatic variables are a key factor in tree interactions in mixed stands of beech (Fagus sylvatica L.) and Scots pine (Pinus sylvestris L.). To test how site climatic conditions modify the effect of inter-specific competition on tree basal area growth and tree mortality, growth and mortality models were fitted using monospecific and mixed sample plots located in matching site conditions selected from the Spanish National Forest Inventory in the Navarra Province. Tree competition status was broken down into four terms according to size-symmetry (size-symmetric vs. size-asymmetric) and species identity (intra-specific vs. inter-specific). The results showed that the size-symmetric inter-specific component was non-significant on beech basal area growth and had a negative effect on pine growth. The effect of size-asymmetric inter-specific competition was always non-significant, resulting in a higher basal area growth when the admixed species are of a larger size. The interaction between annual precipitation and this inter-specific competition effect was much more pronounced on beech than on pine. Inter-specific competition had a negative effect on pine growth under better climatic conditions and a positive effect at dryer sites, while beech always benefited from the presence of pine, although the benefit was greater where climatic site conditions were better. In tree mortality models, pine mortality increased with the proportion of beech, while beech mortality was lower as the proportion of pine increased. Precipitation modified the inter-specific competition effect on tree mortality although the site influence was less relevant than on tree growth. For pine mortality, the negative effect of beech admixture was stronger at lower mean annual precipitation, while in the case of beech the positive effect of pine increased at higher levels of precipitation. The influence of climate on the effect of competition, the variation in their strength depending on the mode of competition (size-symmetric or size-asymmetric), along with the inter-specific competition component, highlight the importance of considering the effect of site conditions on between-species interactions when modeling tree growth and mortality. The species-specific patterns of variation in tree interactions along climatic gradients and the differences in tree growth and mortality corroborate the need to consider the nature of stress-limiting factors and species composition and the importance of analyzing both dynamic processes simultaneously.


Species interaction Abiotic variable Competition reduction Climate–competition interaction Above- and belowground resources Precipitation 



The authors acknowledge Roberto Vallejo, in charge of the Spanish National Forest Inventory, for the provision of the NFI data. They also thank the Spanish Ministry of Economy and Competitiveness for funding the research project “Mixed Forest complexity and sustainability: dynamic, silviculture and adaptive management tools” (AGL2014-51964-C2-2-R).


  1. Adame P, Del Río M, Cañellas I (2010) Modeling individual-tree mortality in Pyrenean oak (Quercus pyrenaica Willd.) stands. Ann For Sci 67:810CrossRefGoogle Scholar
  2. Amoroso MM, Turnblom EC (2006) Comparing productivity of pure and mixed Douglas-fir and western hemlock plantations in the Pacific Northwest. Can J For Res 36:1484–1496CrossRefGoogle Scholar
  3. Assmann E (1970) The principles of yield study. Studies in the organic production, structure, increment and yield of forest stands. Oxford Pergamon Press, OxfordGoogle Scholar
  4. Begon M, Townsend CR, Harper JL (2006) Ecology: from individuals to ecosystems, 4th edn. Blackwell Publishing, MaldenGoogle Scholar
  5. Bertness MD, Callaway R (1994) Positive interactions in communities. Trends Ecol Evol 9:191–193CrossRefPubMedGoogle Scholar
  6. Bravo-Oviedo A, Sterba H, Del Río M, Bravo F (2006) Competition-induced mortality for Mediterranean Pinus pinaster Ait. and P. sylvestris L. For Ecol Manage 222:88–98CrossRefGoogle Scholar
  7. Bravo-Oviedo A, Pretzsch H, Ammer C, Andenmatten E, Antón C, Barbati A, Barreiro S, Brang P, Bravo F, Brunner A, Coll L, Corona M, Den Ouden J, Drössler L, Ducey MJ, Kaynas BY, Legay M, Löf M, Lesinski J, Mason B, Meliadis M, Manetti MC, Morneau F, Motiejunaite J, O’Reilly C, Pach M, Ponette Q, Río M, Short I, Skovsgaard JP, Souidi Z, Spathelf P, Sterba H, Stojanovic D, Strelcova K, Svoboda M, Valsta L, Verheyen K, Zlatanov T (2014) European mixed forests: definition and perspectives. For Syst 23(3):518–533Google Scholar
  8. Canham CD, Uriarte M (2006) Analysis of neighborhood dynamics of forest ecosystems using likelihood methods and modeling. Ecol Appl 16:62–73CrossRefPubMedGoogle Scholar
  9. Canham CD, LePage PT, Coates KD (2004) A neighborhood analysis of canopy tree competition: effects of shading versus crowding. Can J For Res 34:778–787CrossRefGoogle Scholar
  10. Cavard X, Bergeron Y, Chen HY, Paré D, Laganière J, Brassard B (2011) Competition and facilitation between tree species change with stand development. Oikos 120:1683–1695CrossRefGoogle Scholar
  11. Coates KD, Canham CD, LePage PT (2009) Above-versus below-ground competitive effects and responses of a guild of temperate tree species. J Ecol 97:118–130CrossRefGoogle Scholar
  12. Coates KD, Lilles EB, Astrup R (2013) Competitive interactions across a soil fertility gradient in a multispecies forest. J Ecol 101:806–818CrossRefGoogle Scholar
  13. Condés S, Del Rio M, Sterba H (2013) Mixing effect on volume growth of Fagus sylvatica and Pinus sylvestris is modulated by stand density. For Ecol Manage 292:86–95CrossRefGoogle Scholar
  14. Curt T, Prévosto B (2003) Rooting strategy of naturally regenerated beech in Silver birch and Scots pine woodlands. In: Roots: the dynamic interface between plants and the Earth. Springer Netherlands, pp 265–279Google Scholar
  15. Dieler J, Pretzsch H (2013) Morphological plasticity of European beech (Fagus sylvatica L.) in pure and mixed-species stands. For Ecol Manage 295:97–108CrossRefGoogle Scholar
  16. Forrester DI (2014) The spatial and temporal dynamics of species interactions in mixed-species forests: from pattern to process. For Ecol Manage 312:282–292CrossRefGoogle Scholar
  17. Forrester DI, Kohnle U, Albrecht AT, Bauhus J (2013) Complementarity in mixed-species stands of Abies alba and Picea abies varies with climate, site quality and stand density. For Ecol Manage 304:233–242CrossRefGoogle Scholar
  18. Gamfeldt L, Snäll T, Bagchi R, Jonsson M, Gustafsson L, Kjellander P, Ruiz-Jaen MC, Fröberg M, Stendahl J, Philipson CD (2013) Higher levels of multiple ecosystem services are found in forests with more tree species. Nat Commun 4:1340PubMedCentralCrossRefPubMedGoogle Scholar
  19. Garber SM, Maguire DA (2004) Stand productivity and development in two mixed-species spacing trials in the central Oregon Cascades. For Sci 50:92–105Google Scholar
  20. Gómez-Aparicio L, García-Valdés R, Ruiz-Benito P, Zavala MA (2011) Disentangling the relative importance of climate, size and competition on tree growth in Iberian forests: implications for forest management under global change. Global Change Biol 17:2400–2414CrossRefGoogle Scholar
  21. Gonzalo-Jiménez J (2010) Diagnosis fitoclimática de la España Peninsular: hacia un modelo de clasificación funcional de la vegetación y de los ecosistemas peninsulares españoles. Organismo Autónomo de Parques NacionalesGoogle Scholar
  22. Grime JP (1979) Plant strategies and vegetation processes. Wiley, LondonGoogle Scholar
  23. Hugershoff R (1936) Die mathematischen Hilfsmittel der Kulturingenieurs und Biologen, vol 2. Herleitung von gesetzmäßigen Zusammenhängen als Manuskript veröffentlicht, DresdenGoogle Scholar
  24. Jactel H, Nicoll BC, Branco M, Gonzalez-Olabarria JR, Grodzki W, Långström B, Moreira F, Netherer S, Orazio C, Piou D (2009) The influences of forest stand management on biotic and abiotic risks of damage. Ann For Sci 66:1–18CrossRefGoogle Scholar
  25. Johnson PCD (2014) Extension Nakagawa & Schielzeth’s R2 GLMM to random slopes models. Methods Ecol Evol 5:44–946CrossRefGoogle Scholar
  26. Jutras S, Hokka H, Alenius V, Salminen H (2003) Modeling mortality of individual trees in drained peatland sites in Finland. Silva Fenn 37:235–251CrossRefGoogle Scholar
  27. Larocque GR, Luckai N, Adhikary SN, Groot A, Bell FW, Sharma M (2013) Competition theory-science and application in mixed forest stands: review of experimental and modelling methods and suggestions for future research. Environ Rev 21:71–84CrossRefGoogle Scholar
  28. Lebourgeois F, Gomez N, Pinto P, Mérian P (2013) Mixed stands reduce Abies alba tree-ring sensitivity to summer drought in the Vosges mountains, western Europe. For Ecol Manage 303:61–71CrossRefGoogle Scholar
  29. Lines ER, Coomes DA, Purves DW (2010) Influences of forest structure, climate and species composition on tree mortality across the eastern US. PLoS One 5:e13212PubMedCentralCrossRefPubMedGoogle Scholar
  30. Lortie CJ, Callaway RM (2006) Re-analysis of meta-analysis: support for the stress-gradient hypothesis. J Ecol 94:7–16CrossRefGoogle Scholar
  31. Maestre FT, Callaway RM, Valladares F, Lortie CJ (2009) Refining the stress-gradient hypothesis for competition and facilitation in plant communities. J Ecol 97:199–205CrossRefGoogle Scholar
  32. Monserud RA, Sterba H (1999) Modeling individual tree mortality for Austrian forest species. For Ecol Manage 113:109–123CrossRefGoogle Scholar
  33. Nakagawa S, Schielzeth H (2013) A general and simple method for obtaining R2 from Generalized Linear Mixed-effects Models. Methods Ecol Evol 4:133–142CrossRefGoogle Scholar
  34. Perot T, Picard N (2012) Mixture enhances productivity in a two-species forest: evidence from a modeling approach. Ecol Res 27:83–94CrossRefGoogle Scholar
  35. Pretzsch H, Biber P (2005) A re-evaluation of Reineke’s rule and stand density index. For Sci 51:304–320Google Scholar
  36. Pretzsch H, Biber P (2010) Size-symmetric versus size-asymmetric competition and growth partitioning among trees in forest stands along an ecological gradient in central Europe. Can J For Res 40:370–384CrossRefGoogle Scholar
  37. Pretzsch H, Block J, Dieler J, Dong PH, Kohnle U, Nagel J, Spellmann H, Zingg A (2010) Comparison between the productivity of pure and mixed stands of Norway spruce and European beech along an ecological gradient. Ann For Sci 67:1–12CrossRefGoogle Scholar
  38. Pretzsch H, Bielak K, Block J, Bruchwald A, Dieler J, Ehrhart H-P, Kohnle U, Nagel J, Spellmann H, Zasada M (2013a) Productivity of mixed versus pure stands of oak (Quercus petraea (MATT.) LIEBL. and Quercus robur L.) and European beech (Fagus sylvatica L.) along an ecological gradient. Eur J For Res 132(2):263–280CrossRefGoogle Scholar
  39. Pretzsch H, Schütze G, Uhl E (2013b) Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter-specific facilitation. Plant Biol 15:483–495CrossRefPubMedGoogle Scholar
  40. Pretzsch H, Del Río M, Ammer Ch, Avdagic A., Barbeito I, Bielak K, Brazaitis G, Coll L, Dirnberger G, Drössler L, Fabrika M, Forrester DI, Godvod K, Heym M, Hurt V, Kurylyak V, Löf M, Lombardi F, Mohren F, Motta R, den Ouden J, Pach M, Ponette Q, Schütze G, Schweig J, Skrzyszewski J, Sramek V, Sterba H, Stojanović D, Svoboda M, Vanhellemont M, Verheyen K, Wellhausen K, Zlatanov T, Bravo-Oviedo A (2015) Growth and yield of mixed versus pure stands of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvativa L.) analyzed along a productivity gradient through Europe. Eur J For Res 134(5):927–947CrossRefGoogle Scholar
  41. R Core Team (2014) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
  42. Reineke LH (1933) Perfecting a stand-density index for even-aged forests. J Agric Res 46:627–638Google Scholar
  43. Río M, Montero G, Bravo F (2001) Analysis of diameter-density relationships and self-thinning in non-thinned even-aged Scots pine stands. For Ecol Manage 142:79–87CrossRefGoogle Scholar
  44. Río M, Condés S, Pretzsch H (2014) Analyzing size-symmetric vs. size-asymmetric and intra-vs. inter-specific competition in beech (Fagus sylvatica L.) mixed stands. For Ecol Manage 325:90–98CrossRefGoogle Scholar
  45. Rothe A, Binkley D (2001) Nutritional interactions in mixed species forests: a synthesis. Can J For Res 31(11):1855–1870CrossRefGoogle Scholar
  46. Ruiz-Benito P, Lines ER, Gómez-Aparicio L, Zavala MA, Coomes DA (2013) Patterns and drivers of tree mortality in Iberian forests: climatic effects are modified by competition. PLoS One 8:e56843PubMedCentralCrossRefPubMedGoogle Scholar
  47. Sabaté S, Gracia CA, Sánchez A (2002) Likely effects of climate change on growth of Quercus ilex, Pinus halepensis, Pinus pinaster, Pinus sylvestris and Fagus sylvatica forests in the Mediterranean region. For Ecol Manage 162:23–37CrossRefGoogle Scholar
  48. Schwinning S, Weiner J (1998) Mechanisms determining the degree of size asymmetry in competition among plants. Oecologia 113:447–455CrossRefGoogle Scholar
  49. Soliveres S, Maestre FT (2014) Plant–plant interactions, environmental gradients and plant diversity: a global synthesis of community-level studies. Perspect Plant Ecol Evol Syst 16:154–163PubMedCentralCrossRefPubMedGoogle Scholar
  50. Stephenson NL, Van Mantgem PJ, Bunn AG, Bruner H, Harmon ME, O’Connell KB, Urban DL, Franklin JF (2011) Causes and implications of the correlation between forest productivity and tree mortality rates. Ecol Monogr 81:527–555CrossRefGoogle Scholar
  51. Sterba H, Del Rio M, Brunner A, Condes S (2014) Effect of species proportion definition on the evaluation of growth in pure vs. mixed stands. For Syst 23:547–559Google Scholar
  52. Toïgo M, Vallet P, Perot T, Bontemps JD, Piedallu C, Courbaud B (2015) Overyielding in mixed forests decreases with site productivity. J Ecol 103(2):502–512CrossRefGoogle Scholar
  53. Toledo M, Poorter L, Peña-Claros M, Alarcón A, Balcázar J, Leaño C, Licona JC, Llanque O, Vroomans V, Zuidema P (2011) Climate is a stronger driver of tree and forest growth rates than soil and disturbance. J Ecol 99:254–264CrossRefGoogle Scholar
  54. Vilà M, Inchausti P, Vayreda J, Barrantes O, Gracia C, Ibáñez JJ, Mata T (2005) Confounding factors in the observational productivity-diversity relationship in forests. In: Forest diversity and function. Springer Berlin Heidelberg, pp 65-86Google Scholar
  55. Vilà-Cabrera A, Martínez-Vilalta J, Vayreda J, Retana J (2011) Structural and climatic determinants of demographic rates of Scots pine forests across the Iberian Peninsula. Ecol Appl 21:1162–1172CrossRefPubMedGoogle Scholar
  56. Weiner J, Thomas SC (1986) Size variability and competition in plant monocultures. Oikos 47:211–222CrossRefGoogle Scholar
  57. Weiner J, Berntson GM, Thomas SC (1990) Competition and growth form in a woodland annual. J Ecol 78:459-469CrossRefGoogle Scholar
  58. Weiskittel AR, Hann DW, Kershaw Jr JA, Vanclay JK (2011) Forest growth and yield modeling. Wiley, ChichesterGoogle Scholar
  59. Wykoff WR (1990) A basal area increment model for individual conifers in the northern Rocky Mountains. For Sci 36:1077–1104Google Scholar
  60. Yao X, Titus SJ, MacDonald SE (2001) A generalized logistic model of individual tree mortality for aspen, white spruce, and lodgepole pine in Alberta mixedwood forests. Can J For Res 31:283–291Google Scholar
  61. Zhao D, Borders B, Wilson M (2004) Individual-tree diameter growth and mortality models for bottomland mixed-species hardwood stands in the lower Mississippi alluvial valley. For Ecol Manage 199:307–322CrossRefGoogle Scholar
  62. Zhao D, Borders B, Wilson M, Rathbun SL (2006) Modeling neighborhood effects on the growth and survival of individual trees in a natural temperate species-rich forest. Ecol Model 196:90–102CrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2015

Authors and Affiliations

  1. 1.Department of Natural Systems and Resources, School of ForestryTechnical University of MadridMadridSpain
  2. 2.Department of Silviculture and Forest ManagementINIA, Forest Research CentreMadridSpain
  3. 3.Sustainable Forest Management Research InstituteUniversity of Valladolid - INIAMadridSpain

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