Ecological and Physiological Processes in Mixed Versus Monospecific Stands

  • David I. ForresterEmail author


Tree growth depends on the resource availability, the proportion of resources acquired, and the efficiency with which those resources are used. Each of these variables can be influenced by species interactions. These interactions are dynamic and change spatially and temporally as resource availability and climatic conditions change. It is important to understand these processes when designing and managing mixed-species stands and also when modelling these processes. These interactions and their dynamics are the focus of this chapter. To begin with, the production ecology equation is described because it provides a useful framework to quantify the types of processes that influence the growth of forests and how these are influenced by species interactions. This equation describes growth as a function of resource availability, resource acquisition, and resource-use efficiency. Then, while referring to this equation, some of the main types of processes are described in terms of how they influence these variables and hence the productivity of mixtures. This is done for nutrients, then light, and then water. The influence of a given type of interaction on growth is not static. Instead, it changes with spatial and temporal variability in resource availability and climatic conditions and as a stand develops. Therefore, the next section describes a framework that explains these spatial and temporal dynamics and indicates when different types of interactions are important. Finally, stand density can influence the effect of these interactions. As stand density increases, interactions may become more favourable or less favourable, depending on how, and which, resources are influenced by the change in density. The final section therefore shows why stand density needs to be taken into account when examining how species interact.


Stand Density Species Interaction Interspecific Difference Hydraulic Redistribution Complementarity Effect 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


  1. Anten NPR (2005) Optimal photosynthetic characteristics of individual plants in vegetation stands and implications for species coexistence. Ann Bot 95:495–506PubMedCrossRefGoogle Scholar
  2. Anten NPR, Shieving F, Werger MJA (1995) Patterns of light and nitrogen distribution in relation to whole canopy carbon gain in C3 and C4 mono- and dicotyledonous species. Oecologia 101:504–513PubMedCrossRefGoogle Scholar
  3. Augusto L, Ranger J, Binkley D, Rothe A (2002) Impact of several common tree species of European temperate forests on soil fertility. Ann For Sci 59:233–253CrossRefGoogle Scholar
  4. Augusto L, Schrijver AD, Vesterdal L, Smolander A, Prescott C, Ranger J (2015) Influences of evergreen gymnosperm and deciduous angiosperm tree species on the functioning of temperate and boreal forests. Biol Rev 90:444–466PubMedCrossRefGoogle Scholar
  5. Bauhus J, Khanna PK, Menden N (2000) Aboveground and belowground interactions in mixed plantations of Eucalyptus globulus and Acacia mearnsii. Can J For Res 30:1886–1894CrossRefGoogle Scholar
  6. Bauhus J, van Winden AP, Nicotra AB (2004) Above-ground interactions and productivity in mixed-species plantations of Acacia mearnsii and Eucalyptus globulus. Can J For Res 34:686–694CrossRefGoogle Scholar
  7. Bayer D, Seifert S, Pretzsch H (2013) Structural crown properties of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) in mixed versus pure stands revealed by terrestrial laser scanning. Trees Struct Funct 27:1035–1047CrossRefGoogle Scholar
  8. Berger TW, Inselsbacher E, Mutsch F, Pfeffer M (2009a) Nutrient cycling and soil leaching in eighteen pure and mixed stands of beech (Fagus sylvatica) and spruce (Picea abies). For Ecol Manag 258:2578–2592CrossRefGoogle Scholar
  9. Berger TW, Untersteiner H, Toplitzer M, Neubauer C (2009b) Nutrient fluxes in pure and mixed stands of spruce (Picea abies) and beech (Fagus sylvatica). Plant Soil 322:317–342CrossRefGoogle Scholar
  10. Binkley D (2003) Seven decades of stand development in mixed and pure stands of conifers and nitrogen-fixing red alder. Can J For Res 33:2274–2279CrossRefGoogle Scholar
  11. Binkley D (2012) Understanding the role resource use efficiency in determining the growth of trees and forests. In: Schlichter T, Montes L (eds) Forests in development: A vital balance. Springer, The Hague, pp 13–26Google Scholar
  12. Binkley D, Giardina C (1997) Nitrogen fixation in tropical forest plantations. In: Nambiar EKS, Brown AG (eds) Management of soil, nutrients and water in tropical plantation forests. Australian Centre for International Agricultural Research, Canberra, pp 297–337Google Scholar
  13. Binkley D, Giardina C (1998) Why do tree species affect soils? The Warp and Woof of tree-soil interactions. Biogeochemistry 42:89–106CrossRefGoogle Scholar
  14. Binkley D, Valentine D (1991) Fifty-year biogeochemical effects of green ash, white pine, and Norway spruce in a replicated experiment. For Ecol Manag 40(1–2):13–25CrossRefGoogle Scholar
  15. Binkley D, Dunkin KA, DeBell D, Ryan MG (1992a) Production and nutrient cycling in mixed plantations of Eucalyptus and Albizia in Hawaii. For Sci 38(2):393–408Google Scholar
  16. Binkley D, Sollins P, Bell R, Sachs D, Myrold D (1992b) Biogeochemistry of adjacent conifer and alder-conifer stands. Ecology 73:2022–2033CrossRefGoogle Scholar
  17. Binkley D, Senock R, Bird S, Cole TG (2003) Twenty years of stand development in pure and mixed stands of Eucalyptus saligna and N-fixing Facaltaria moluccana. For Ecol Manag 182:93–102CrossRefGoogle Scholar
  18. Binkley D, Stape JL, Ryan MG (2004) Thinking about efficiency of resource use in forests. For Ecol Manag 193:5–16CrossRefGoogle Scholar
  19. Bolte A, Villanueva I (2006) Interspecific competition impacts on the morphology and distribution of fine roots in European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.) Eur J For Res 125:15–26CrossRefGoogle Scholar
  20. Bouillet JP, Laclau JP, Goncalves JLM, Moreira MZ, Trivelin PCO, Jourdan C, Silva EV, Piccolo MC, Tsai SM, Galiana A (2008) Mixed-species plantations of Acacia mangium and Eucalyptus grandis in Brazil 2: nitrogen accumulation in the stands and biological N2 fixation. For Ecol Manag 255:3918–3930CrossRefGoogle Scholar
  21. Bouillet J-P, Laclau J-P, Gonçalves JLM, Voigtlaender M, Gava JL, Leite FP, Hakamada R, Mareschal L, Mabiala A, Tardy F, Levillain J, Deleporte P, Epron D, Nouvellon Y (2013) Eucalyptus and Acacia tree growth over entire rotation in single- and mixed-species plantations across five sites in Brazil and Congo. For Ecol Manag 301:89–101CrossRefGoogle Scholar
  22. Boyden S, Binkley D, Senock R (2005) Competition and facilitation between Eucalyptus and nitrogen-fixing Falcataria in relation to soil fertility. Ecology 86(4):992–1001CrossRefGoogle Scholar
  23. Boyden S, Montgomery R, Reich PB, Palik B (2012) Seeing the forest for the heterogeneous trees: stand-scale resource distributions emerge from tree-scale structure. Ecol Appl 22(5):1578–1588PubMedCrossRefGoogle Scholar
  24. Brassard BW, Chen HYH, Bergeron Y, Paré D (2011) Differences in fine root productivity between mixed- and single-species stands. Funct Ecol 25:238–246CrossRefGoogle Scholar
  25. Brassard BW, Chen HYH, Cavard X, Laganière J, Reich PB, Bergeron Y, Paré D, Yuan Z (2013) Tree species diversity increases fine root productivity through increased soil volume filling. J Ecol 101:210–219CrossRefGoogle Scholar
  26. Canham C, Coates KD, Bartemucci P, Quaglia S (1999) Measurement and modeling of spatially explicit variation in light transmission through interior cedar-hemlock forests of British Columbia. Can J For Res 29:1775–1783CrossRefGoogle Scholar
  27. Cavard X, Bergeron Y, Chen HYH, Paré D, Laganière J, Brassard B (2011) Competition and facilitation between tree species change with stand development. Oikos 120:1683–1695CrossRefGoogle Scholar
  28. Chisholm RA, Muller-Landau HC, Rahman KA, Bebber DP, Bin Y, Bohlman SA, Bourg NA, Brinks J, Bunyavejchewin S, Butt N, Cao H, Cao M, Cardenas D, Chang L-W, Chiang J-M, Chuyong G, Condit R, Dattaraja HS, Davies S, Duque A, Fletcher C, Gunatilleke N, Gunatilleke S, Hao Z, Harrison RD, Howe R, Hsieh C-F, Hubbell SP, Itoh A, Kenfack D, Kiratiprayoon S, Larson AJ, Lian J, Lin D, Liu H, Lutz JA, Ma K, Malhi Y, McMahon S, McShea W, Meegaskumbura M, Razman SM, Morecroft MD, Nytch CJ, Oliveira A, Parker GG, Pulla S, Punchi-Manage R, Romero-Saltos H, Sang W, Schurman J, Su S-H, Sukumar R, Sun I-F, Suresh HS, Tan S, Thomas D, Thomas S, Thompson J, Valencia R, Wolf A, Yap S, Ye W, Yuan Z, Zimmerman JK (2013) Scale-dependent relationships between tree species richness and ecosystem function in forests. J Ecol 101:1214–1224CrossRefGoogle Scholar
  29. Coates KD, Lilles EB, Astrup R (2013) Competitive interactions across a soil fertility gradient in a multispecies forest. J Ecol 101(3):806–818CrossRefGoogle Scholar
  30. Condés S, Rio MD, Sterba H (2013) Mixing effect on volume growth of Fagus sylvatica and Pinus sylvestris is modulated by stand density. For Ecol Manag 292:86–95CrossRefGoogle Scholar
  31. Crews TE, Peoples MB (2005) Can the synchrony of nitrogen supply and crop demand be improved in legume and fertilizer-based agroecosystems? A review. Nutr Cycl Agroecosyst 72:101–120CrossRefGoogle Scholar
  32. DeBell DS, Radwan MA (1979) Growth and nitrogen relations of coppiced black cottonwood and red alder in pure and mixed plantations. Bot Gaz 140(Suppl.):S97–S101Google Scholar
  33. DeBell DS, Whitesell CD, Crabb TB (1987) Benefits of Eucalyptus-Albizia mixtures vary by site on Hawaii Island. USDA For. Serv. Res. Paper PSW-187Google Scholar
  34. del Río M, Sterba H (2009) Comparing volume growth in pure and mixed stands of Pinus sylvestris and Quercus pyrenaica. Ann For Sci 66:502CrossRefGoogle Scholar
  35. del Río M, Schütze G, Pretzsch H (2014) Temporal variation of competition and facilitation in mixed species forests in Central Europe. Plant Biol 16:166–176PubMedCrossRefGoogle Scholar
  36. Dewar RC, Tarvainen L, Parker K, Wallin G, McMurtrie RE (2012) Why does leaf nitrogen decline within tree canopies less rapidly than light? An explanation from optimization subject to a lower bound on leaf mass per area. Tree Physiol 32:520–534PubMedCrossRefGoogle Scholar
  37. Dieler J, Pretzsch H (2013) Morphological plasticity of European beech (Fagus sylvatica L.) in pure and mixed-species stands. For Ecol Manag 295:97–108CrossRefGoogle Scholar
  38. Dijkstra FA, West JB, Hobbie SE, Reich PB (2009) Antagonistic effects of species on C respiration and net N mineralization in soils from mixed coniferous plantations. For Ecol Manag 257:1112–1118CrossRefGoogle Scholar
  39. Dunn GM, Connor DJ (1993) An analysis of sap flow in Mountain Ash (Eucalyptus regnans) forests of different age. Tree Physiol 13:321–336PubMedCrossRefGoogle Scholar
  40. Epron D, Nouvellon Y, Mareschal L, MoreiraeMoreira R, Koutika L-S, Geneste B, Delgado-Rojas JS, Laclau J-P, Sola G, Gonçalves JLM, Bouillet J-P (2013) Partitioning of net primary production in Eucalyptus and Acacia stands and in mixed-species plantations: two case-studies in contrasting tropical environments. For Ecol Manag 301:102–111CrossRefGoogle Scholar
  41. Ewel JJ (1986) Designing agricultural ecosystems for the humid tropics. Annu Rev Ecol Syst 17:245–271CrossRefGoogle Scholar
  42. Field C (1983) Allocating leaf nitrogen for the maximization of carbon gain: leaf age as a control on the allocation program. Oecologia 56:341–347PubMedCrossRefGoogle Scholar
  43. Filipescu CN, Comeau PG (2007) Competitive interactions between aspen and white spruce vary with stand age in boreal mixedwoods. For Ecol Manag 247:175–184CrossRefGoogle Scholar
  44. Fisher RF, Binkley D (2000) Ecology and management of forest soils, 3rd edn. Wiley, New York, ChichesterGoogle Scholar
  45. Forrester DI (2013) Growth responses to thinning, pruning and fertiliser application in Eucalyptus plantations: a review of their production ecology and interactions. For Ecol Manag 310:336–347CrossRefGoogle Scholar
  46. Forrester DI (2014) The spatial and temporal dynamics of species interactions in mixed-species forests: from pattern to process. For Ecol Manag 312:282–292CrossRefGoogle Scholar
  47. Forrester DI (2015) Transpiration and water-use efficiency in mixed-species forests versus monocultures: effects of tree size, stand density and season. Tree Physiol 35:289–304PubMedCrossRefGoogle Scholar
  48. Forrester DI, Albrecht AT (2014) Light absorption and light-use efficiency in mixtures of Abies alba and Picea abies along a productivity gradient. For Ecol Manag 328:94–102CrossRefGoogle Scholar
  49. Forrester DI, Bauhus J (2016) A review of processes behind diversity – productivity relationships in forests. Curr Forest Rep 2:45–61CrossRefGoogle Scholar
  50. Forrester DI, Pretzsch H (2015) Tamm Review: on the strength of evidence when comparing ecosystem functions of mixtures with monocultures. For Ecol Manag 356:41–53CrossRefGoogle Scholar
  51. Forrester DI, Smith RGB (2012) Faster growth of Eucalyptus grandis and Eucalyptus pilularis in mixed-species stands than monocultures. For Ecol Manag 286:81–86CrossRefGoogle Scholar
  52. Forrester DI, Bauhus J, Cowie AL (2005) Nutrient cycling in a mixed-species plantation of Eucalyptus globulus and Acacia mearnsii. Can J For Res 35(12):2942–2950. doi: 10.1139/x05-214 CrossRefGoogle Scholar
  53. Forrester DI, Bauhus J, Cowie AL (2006a) Carbon allocation in a mixed-species plantation of Eucalyptus globulus and Acacia mearnsii. For Ecol Manag 233:275–284. doi: 10.1016/j.foreco.2006.05.018 CrossRefGoogle Scholar
  54. Forrester DI, Bauhus J, Cowie AL, Vanclay JK (2006b) Mixed-species plantations of Eucalyptus with nitrogen fixing trees: a review. For Ecol Manag 233:211–230. doi: 10.1016/j.foreco.2006.05.012 CrossRefGoogle Scholar
  55. Forrester DI, Cowie AL, Bauhus J, Wood J, Forrester RI (2006c) Effects of changing the supply of nitrogen and phosphorus on growth and interactions between Eucalyptus globulus and Acacia mearnsii in a pot trial. Plant Soil 280(1-2):267–277. doi: 10.1007/s11104-005-3228-x CrossRefGoogle Scholar
  56. Forrester DI, Bauhus J, Cowie AL, Mitchell PA, Brockwell J (2007a) Productivity of three young mixed-species plantations containing N2-fixing Acacia and non-N2-fixing Eucalyptus and Pinus trees in Southeastern Australia. For Sci 53(3):426–434Google Scholar
  57. Forrester DI, Schortemeyer M, Stock WD, Bauhus J, Khanna PK, Cowie AL (2007b) Assessing nitrogen fixation in mixed- and single-species plantations of Eucalyptus globulus and Acacia mearnsii. Tree Physiol 27:1319–1328. doi: 10.1093/treephys/27.9.1319 PubMedCrossRefGoogle Scholar
  58. Forrester DI, Collopy JJ, Morris JD (2010a) Transpiration along an age series of Eucalyptus globulus plantations in southeastern Australia. For Ecol Manag 259:1754–1760. doi: 10.1016/j.foreco.2009.04.023 CrossRefGoogle Scholar
  59. Forrester DI, Theiveyanathan S, Collopy JJ, Marcar NE (2010b) Enhanced water use efficiency in a mixed Eucalyptus globulus and Acacia mearnsii plantation. For Ecol Manag 259:1761–1770. doi: 10.1016/j.foreco.2009.07.036 CrossRefGoogle Scholar
  60. Forrester DI, Vanclay JK, Forrester RI (2011) The balance between facilitation and competition in mixtures of Eucalyptus and Acacia changes as stands develop. Oecologia 166(1):265–272PubMedCrossRefGoogle Scholar
  61. Forrester DI, Collopy JJ, Beadle CL, Baker TG (2012a) Interactive effects of simultaneously applied thinning, pruning and fertiliser application treatments on growth, biomass production and crown architecture in a young Eucalyptus nitens plantation. For Ecol Manag 267:104–116. doi: 10.1016/j.foreco.2011.11.039 CrossRefGoogle Scholar
  62. Forrester DI, Lancaster K, Collopy JJ, Warren CR, Tausz M (2012b) Photosynthetic capacity of Eucalyptus globulus is higher when grown in mixture with Acacia mearnsii. Trees Struct Funct 26:1203–1213CrossRefGoogle Scholar
  63. 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 Manag 304:233–242CrossRefGoogle Scholar
  64. Forrester DI, Benneter A, Bouriaud O, Bauhus J (2016) Diversity and competition influence tree allometry – developing allometric functions for mixed-species forests. J Ecol. doi: 10.1111/1365-2745.12704 Google Scholar
  65. Forrester DI, Ammer C, Annighöfer PJ, Barbeito I, Bielak K, Bravo-Oviedo A, Coll L, del Río M, Drössler L, Heym M, Hurt V, Löf M, den Ouden J, Pach M, Pereira MG, Plaga B, Ponette Q, Skrzyszewski J, Sterba H, Svoboda M, Zlatanov T, Pretzsch H (in press) Effects of crown architecture and stand structure on light absorption in mixed and monospecific Fagus sylvatica and Pinus sylvestris forests along a productivity and climate gradient through Europe. J EcolGoogle Scholar
  66. Fredericksen TS, Zedaker SM (1995) Fine root biomass, distribution, and production in young pine-hardwood stands. New For 10:99–110Google Scholar
  67. 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
  68. Gartner TB, Cardon ZG (2004) Decomposition dynamics in mixed-species leaf litter. Oikos 104:230–246CrossRefGoogle Scholar
  69. Gash JHC, Lloyd CR, Lachaud G (1995) Estimating sparse forest rainfall interception with an analytical model. J Hydrol 170:79–86CrossRefGoogle Scholar
  70. Gebauer T, Horna V, Leuschner C (2012) Canopy transpiration of pure and mixed forest stands with variable abundance of European beech. J Hydrol 442–443:2–14CrossRefGoogle Scholar
  71. Gerrits AMJ, Pfister L, Savenije HHG (2010) Spatial and temporal variability of canopy and forest floor interception in a beech forest. Hydrol Process 24:3011–3025CrossRefGoogle Scholar
  72. Getzin S, Wiegand K (2007) Asymmetric tree growth at the stand level: random crown patterns and the response to slope. For Ecol Manag 242:165–174CrossRefGoogle Scholar
  73. Grossiord C, Granier A, Ratcliffe S, Bouriaud O, Bruelheide H, Chećko E, Forrester DI, Dawud SM, Finér L, Pollastrini M, Scherer-Lorenzen M, Valladares F, Bonal D, Gessler A (2014) Tree diversity does not always improve resistance of forest ecosystems to drought. Proc Natl Acad Sci U S A 111(41):14812–14815PubMedPubMedCentralCrossRefGoogle Scholar
  74. Guisasola R, Tang X, Bauhus J, Forrester DI (2015) Intra- and inter-specific differences in crown architecture in Chinese subtropical mixed-species forests. For Ecol Manag 353:164–172CrossRefGoogle Scholar
  75. Harrington RA, Fownes JH, Meinzer FC, Scowcroft PG (1995) Forest growth along a rainfall gradient in Hawaii: Acacia koa stand structure, productivity, foliar nutrients, and water- and nutrient-use efficiencies. Oecologia 102:277–284PubMedCrossRefGoogle Scholar
  76. Hättenschwiler S, Tiunov AV, Scheu S (2005) Biodiversity and litter decomposition in terrestrial ecosystems. Annu Rev Ecol Evol Syst 36:191–218CrossRefGoogle Scholar
  77. Hawthorne SND, Lane PNJ, Bren LJ, Sims NC (2013) The long term effects of thinning treatments on vegetation structure and water yield. For Ecol Manag 310:983–993CrossRefGoogle Scholar
  78. He X-H, Critchley C, Bledsoe C (2003) Nitrogen transfer within and between plants through common mycorrhizal networks (CMNs). Crit Rev Plant Sci 22(6):531–567CrossRefGoogle Scholar
  79. He Z-B, Yang J-J, Du J, Zhao W-Z, Liu H, Chang X-X (2014) Spatial variability of canopy interception in a spruce forest of the semiarid mountain regions of China. Agric For Meteorol 188:58–63CrossRefGoogle Scholar
  80. Hector A, Bagchi R (2007) Biodiversity and ecosystem multifunctionality. Nat Lett 448:188–191CrossRefGoogle Scholar
  81. Hinsinger P, Betencourt E, Bernard L, Brauman A, Plassard C, Shen J, Tang X, Zhang F (2011) P for two, sharing a scarce resource: soil phosphorus acquisition in the rhizosphere of intercropped species. Plant Physiol 156:1078–1086PubMedPubMedCentralCrossRefGoogle Scholar
  82. Hirose T, Werger MJA (1987) Maximizing daily canopy photosynthesis with respect to the leaf nitrogen allocation pattern in the canopy. Oecologia 72(4):520–526PubMedCrossRefGoogle Scholar
  83. Huber MO, Sterba H, Bernhard L (2014) Site conditions and definition of compositional proportion modify mixture effects in Picea abies – Abies alba stands. Can J For Res 44:1281–1291CrossRefGoogle Scholar
  84. Hunt MA, Unwin GL, Beadle CL (1999) Effects of naturally regenerated Acacia dealbata on the productivity of a Eucalyptus nitens plantation in Tasmania, Australia. For Ecol Manag 117:75–85CrossRefGoogle Scholar
  85. Ilek A, Kucza J, Szostek M (2015) The effect of stand species composition on water storage capacity of the organic layers of forest soils. Eur J For Res 134:187–197CrossRefGoogle Scholar
  86. Isbell F, Calcagno V, Hector A, Connolly J, Harpole WS, Reich PB, Scherer-Lorenzen M, Schmid B, Tilman D, Ruijven J, Weigelt A, Wilsey BJ, Zavaleta ES, Loreau M (2011) High plant diversity is needed to maintain ecosystem services. Nature 477:199–203PubMedCrossRefGoogle Scholar
  87. Kaye JP, Resh SC, Kaye MW, Chimmer RA (2000) Nutrient and carbon dynamics in a replacement series of Eucalyptus and Albizia trees. Ecology 81(12):3267–3273CrossRefGoogle Scholar
  88. Kelty MJ (1992) Comparative productivity of monocultures and mixed-species stands. In: Kelty MJ, Larson BC, Oliver CD (eds) The ecology and silviculture of mixed-species forests. Kluwer Academic Publishers, Dordrecht, pp 125–141CrossRefGoogle Scholar
  89. Kelty MJ, Cameron IR (1995) Plot designs for the analysis of species interactions in mixed stands. Commonwealth For Rev 74(4):322–332Google Scholar
  90. Khamzina A, Lamers JPA, Vlek PLG (2009) Nitrogen fixation by Elaeagnus angustifolia in the reclamation of degraded croplands of Central Asia. Tree Physiol 29(6):799–808PubMedCrossRefGoogle Scholar
  91. Khanna PK (1997) Comparison of growth and nutrition of young monocultures and mixed stands of Eucalyptus globulus and Acacia mearnsii. For Ecol Manag 94:105–113CrossRefGoogle Scholar
  92. Khanna PK (1998) Nutrient cycling under mixed-species tree systems in southeast Asia. Agrofor Syst 38:99–120CrossRefGoogle Scholar
  93. Knops JMH, Bradley KL, Wedin DA (2002) Mechanisms of plant species impacts on ecosystem nitrogen cycling. Ecol Lett 5:454–466CrossRefGoogle Scholar
  94. Kranabetter JM, MacKenzie WH (2010) Contrasts among mycorrhizal plant guilds in foliar nitrogen concentration and δ15N along productivity gradients of a boreal forest. Ecosystems 13:108–117CrossRefGoogle Scholar
  95. Kunert N, Schwendenmann L, Potvin C, Hölscher D (2012) Tree diversity enhances tree transpiration in a Panamanian forest plantation. J Appl Ecol 49:135–144CrossRefGoogle Scholar
  96. Laclau J-P, Nouvellon Y, Reine C, Gonçalves JLM, Krushe AV, Jourdan C, le Maire G, Bouillet J-P (2013) Mixing Eucalyptus and Acacia trees leads to fine root over-yielding and vertical segregation between species. Oecologia 172:903–913PubMedCrossRefGoogle Scholar
  97. Law BE, Falge E, Guc L, Baldocchi DD, Bakwind P, Berbigier P, Davis K, Dolmang AJ, Falk M, Fuentes JD, Goldstein A, Granier A, Grelle A, Hollinger D, Janssensm IA, Jarvis P, Jensen NO, Katul G, Mahli Y, Matteucci G, Meyers T, Monsont R, Munger W, Oechel W, Olson R, Pilegaard K, Paw KT, Thorgeirsson H, Valentini R, Verma S, Vesala T, Wilson K, Wofsy S (2002) Environmental controls over carbon dioxide and water vapor exchange of terrestrial vegetation. Agric For Meteorol 113:97–120CrossRefGoogle Scholar
  98. le Maire G, Nouvellon Y, Christina M, Ponzoni FJ, Gonçalves JLM, Bouillet J-P, Laclau J-P (2013) Tree and stand light use efficiencies over a full rotation of single- and mixed-species Eucalyptus grandis and Acacia mangium plantations. For Ecol Manag 288:31–42CrossRefGoogle Scholar
  99. 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 Manag 303:61–71CrossRefGoogle Scholar
  100. Levia DF, Frost EE (2003) A review and evaluation of stemflow literature in the hydrologic and biogeochemical cycles of forested and agricultural ecosystems. J Hydrol 274(1–4):1–29CrossRefGoogle Scholar
  101. Ligot G, Balandier P, Courbaud B, Claessens H (2014) Forest radiative transfer models: which approach for which application? Can J For Res 44(5):385–397CrossRefGoogle Scholar
  102. Litton CM, Raich JW, Ryan MG (2007) Carbon allocation in forest ecosystems. Glob Chang Biol 13:2089–2109CrossRefGoogle Scholar
  103. May BM, Attiwill P, M. (2003) Nitrogen-fixation by Acacia dealbata and changes in soil properties 5 years after mechanical disturbance or slash-burning following timber harvest. For Ecol Manag 181:339-355Google Scholar
  104. McKane RB, Johnson LC, Shaver GR, Nadelhoffer KJ, Rastetter EB, Fry B, Giblin AE, Kielland K, Kwiatkowski BL, Laundre JA, Murray G (2002) Resource-based niches provide a basis for plant species diversity and dominance in arctic tundra. Nature 415:68–71PubMedCrossRefGoogle Scholar
  105. Monteith JL (1977) Climate and the efficiency of crop production in Britain. Philos Trans R Soc Lond B 281:277–294CrossRefGoogle Scholar
  106. Moore GW, Bond BJ, Jones JA (2011) A comparison of annual transpiration and productivity in monoculture and mixed-species Douglas-fir and red alder stands. For Ecol Manag 262:2263–2270CrossRefGoogle Scholar
  107. Neumann RB, Cardon ZG (2012) The magnitude of hydraulic redistribution by plant roots: a review and synthesis of empirical and modeling studies. New Phytol 194:337–352PubMedCrossRefGoogle Scholar
  108. Niinemets Ü (2012) Optimization of foliage photosynthetic capacity in tree canopies: towards identifying missing constraints. Tree Physiol 32:505–509PubMedCrossRefGoogle Scholar
  109. Osada N, Yasumura Y, Ishida A (2014) Leaf nitrogen distribution in relation to crown architecture in the tall canopy species, Fagus crenata. Oecologia 175:1093–1106PubMedCrossRefGoogle Scholar
  110. Parrotta JA, Baker DD, Fried M (1996) Changes in dinitrogen fixation in maturing stands of Casuarina equisetifolia and Leucaena leucocephala. Can J For Res 26(9):1684–1691CrossRefGoogle Scholar
  111. Peltoniemi MS, Duursma RA, Medlyn BE (2012) Co-optimal distribution of leaf nitrogen and hydraulic conductance in plant canopies. Tree Physiol 32:510–519PubMedCrossRefGoogle Scholar
  112. Peoples MB, Herridge DF, Ladha JK (1995) Biological nitrogen fixation: an efficient source of nitrogen for sustainable agricultural production? Plant Soil 174(1-2):3–28CrossRefGoogle Scholar
  113. Pfautsch S, Gessler A, Adams MA, Rennenberg H (2009a) Using amino-nitrogen pools and fluxes to identify contributions of understory Acacia spp. to overstory Eucalyptus regnans and stand nitrogen uptake in temperate Australia. New Phytol 183:1097–1113PubMedCrossRefGoogle Scholar
  114. Pfautsch S, Rennenberg H, Bell TL, Adams MA (2009b) Nitrogen uptake by Eucalyptus regnans and Acacia spp. – preferences, resource overlap and energetic costs. Tree Physiol 29:389–399PubMedCrossRefGoogle Scholar
  115. Poorter H, Niklas KJ, Reich PB, Oleksyn J, Poot P, Mommer L (2012) Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control. New Phytol 193:30–50PubMedCrossRefGoogle Scholar
  116. 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 76:712–723CrossRefGoogle Scholar
  117. Pretzsch H, Bielak K, Block J, Bruchwald A, Dieler J, Ehrhart H-P, Kohnle U, Nagel J, Spellmann H, Zasada M, Zingg A (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 132:263–280CrossRefGoogle Scholar
  118. 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 15:483–495PubMedCrossRefGoogle Scholar
  119. Prieto I, Armas C, Pugnaire FI (2012) Water release through plant roots: new insights into its consequences at the plant and ecosystem level. New Phytol 193:830–841PubMedCrossRefGoogle Scholar
  120. Rao MR, Nair PKR, Ong CK (1998) Biophysical interactions in tropical agroforestry systems. Agrofor Syst 38:3–50CrossRefGoogle Scholar
  121. Reich PB, Luo Y, Bradford JB, Poorter H, Perry CH, Oleksyn J (2014) Temperature drives global patterns in forest biomass distribution in leaves, stems, and roots. Proc Natl Acad Sci U S A 111:13721–13726PubMedPubMedCentralCrossRefGoogle Scholar
  122. Reif A, Brucker U, Kratzer R, Schmiedinger A, Bauhus J (2010) Waldbewirtschaftung in Zeiten des Klimawandels – Synergien und Konfliktpotenziale zwischen Forstwirtschaft und Naturschutz. Naturschutz und Landschaftsplanung 42(9):261–266Google Scholar
  123. Richards AE, Forrester DI, Bauhus J, Scherer-Lorenzen M (2010) The influence of mixed tree plantations on the nutrition of individual species: a review. Tree Physiol 30:1192–1208. doi: 10.1093/treephys/tpq035 PubMedCrossRefGoogle Scholar
  124. Rothe A, Binkley D (2001) Nutritional interactions in mixed species forests: a synthesis. Can J For Res 31(11):1855–1870CrossRefGoogle Scholar
  125. Roupsard O, Ferhi A, Granier A, Pallo F, Depommier D, Mallet B, Joly HI, Dreyer E (1999) Reverse phenology and dry-season water uptake by Faidherbia albida (Del.) A. Chev. in an agroforestry parkland of Sudanese west Africa. Funct Ecol 13:460–472CrossRefGoogle Scholar
  126. Ryan MG, Binkley D, Fownes JH (1997) Age-related decline in forest productivity: pattern and process. Adv Ecol Res 27:213–262CrossRefGoogle Scholar
  127. Ryan MG, Binkley D, Fownes JH, Giardina CP, Senock RS (2004) An experimental test of the causes of forest growth decline with stand age. Ecol Monogr 74:393–414CrossRefGoogle Scholar
  128. Saccone P, Delzon S, Pagès J-P, Brun J-J, Michalet R (2009) The role of biotic interactions in altering tree seedling responses to an extreme climatic event. J Veg Sci 20:403–414CrossRefGoogle Scholar
  129. Sands PJ (1995) Modelling canopy production. I Optimal distribution of photosynthetic resources. Aust J Plant Physiol 22:593–601CrossRefGoogle Scholar
  130. Sapijanskas J, Paquette A, Potvin C, Kunert N, Loreau M (2014) Tropical tree diversity enhances light capture through crown plasticity and spatial and temporal niche differences. Ecology 95:2479–2492CrossRefGoogle Scholar
  131. Schmid I, Kazda M (2002) Root distribution of Norway spruce in monospecific and mixed stands on different soils. For Ecol Manag 159(1-2):37–47CrossRefGoogle Scholar
  132. Schume H, Jost G, Hager H (2004) Soil water depletion and recharge patterns in mixed and pure forest stands of European beech and Norway spruce. J Hydrol 289:258–274CrossRefGoogle Scholar
  133. Schwendenmann L, Pendall E, Sanchez-Bragado R, Kunert N, Hölscher D (2015) Tree water uptake in a tropical plantation varying in tree diversity: interspecific differences, seasonal shifts and complementarity. Ecohydrology 8:1–12CrossRefGoogle Scholar
  134. Snowdon P, Wichiennopparat W, Khanna PK (2003) Growth, above-ground biomass and nutrient content of eucalypts and acacias grown in mixture in a tropical environment – evaluation for one full rotation. In: International Conference on Eucalypt Productivity, Hobart, Australia, 10–15 November 2003. pp 57–58Google Scholar
  135. Stape JL, Binkley D, Ryan MG (2004) Eucalyptus production and the supply, use and efficiency of use of water, light and nitrogen across a geographic gradient in Brazil. For Ecol Manag 193:17–31CrossRefGoogle Scholar
  136. Turner BL (2008) Resource partitioning for soil phosphorus: a hypothesis. J Ecol 96:698–702CrossRefGoogle Scholar
  137. Van Kessel C, Farrell RE, Roskoski JP, Keane KM (1994) Recycling of the naturally-occurring 15N in an established stand of Leucaena leucocephala. Soil Biol Biochem 26:757–762CrossRefGoogle Scholar
  138. Vanclay JK (2006a) Experiment designs to evaluate inter- and intra-specific interactions in mixed plantings of forest trees. For Ecol Manag 233:366–374CrossRefGoogle Scholar
  139. Vanclay JK (2006b) Spatially-explicit competition indices and the analysis of mixed-species plantings with the Simile modelling environment. For Ecol Manag 233:295–302CrossRefGoogle Scholar
  140. Vandermeer J (1989) The ecology of intercropping. Cambridge University Press, New YorkCrossRefGoogle Scholar
  141. Vertessy RA, Hatton TJ, Benyon RG, Dawes WR (1996) Long-term growth and water balance predictions for a mountain ash (Eucalyptus regnans) forest catchment subject to clear-felling and regeneration. Tree Physiol 16(1-2):221–232PubMedCrossRefGoogle Scholar
  142. Vilà M, Carrillo-Gavilán A, Vayreda J, Bugmann H, Fridman J, Grodzki W, Haase J, Kunstler G, Schelhaas M, Trasobares A (2013) Disentangling biodiversity and climatic determinants of wood production. PLoS One 8(2):e53530PubMedPubMedCentralCrossRefGoogle Scholar
  143. Vitousek PM (1982) Nutrient cycling and nutrient efficiency. Am Nat 119:553–572CrossRefGoogle Scholar
  144. Wacker L, Baudois O, Eichenberger-Glinz S, Schmid B (2009) Effects of plant species richness on stand structure and productivity. J Plant Ecol 2(2):95–106CrossRefGoogle Scholar
  145. Wang XL, Klinka K, Chen HYH, de Montigny L (2002) Root structure of western hemlock and western redcedar in single- and mixed-species stands. Can J For Res 32:997–1004CrossRefGoogle Scholar
  146. Williams BL (1992) Nitrogen dynamics in humus and soil beneath Sitka spruce (Picea sitchensis (Bong.) Carr.) planted in pure stands and in mixture with Scots pine (Pinus sylvestris L.) Plant Soil 144:77–84CrossRefGoogle Scholar
  147. Zapater M, Hossann C, Bréda N, Bréchet C, Bonal D, Granier A (2011) Evidence of hydraulic lift in a young beech and oak mixed forest using 18O soil water labelling. Trees 25:885–894CrossRefGoogle Scholar

Copyright information

© Springer-Verlag GmbH Germany 2017

Authors and Affiliations

  1. 1.Swiss Federal Institute for Forest, Snow and Landscape Research WSLBirmensdorfSwitzerland

Personalised recommendations