New Forests

, Volume 43, Issue 5–6, pp 755–770 | Cite as

Increase in size and nitrogen concentration enhances seedling survival in Mediterranean plantations. Insights from an ecophysiological conceptual model of plant survival

  • Pedro Villar-SalvadorEmail author
  • Jaime Puértolas
  • Bárbara Cuesta
  • Juan L. Peñuelas
  • Mercedes Uscola
  • Norberto Heredia-Guerrero
  • José M. Rey Benayas


Reduction in size and tissue nutrient concentration is widely considered to increase seedling drought resistance in dry and oligotrophic plantation sites. However, much evidence indicates that increase in size and tissue nutrient concentration improves seedling survival in Mediterranean forest plantations. This suggests that the ecophysiological processes and functional attributes relevant for early seedling survival in Mediterranean climate must be reconsidered. We propose a ecophysiological conceptual model for seedling survival in Mediterranean-climate plantations to provide a physiological explanation of the frequent positive relationship between outplanting performance and seedling size and nutrient concentration. The model considers the physiological processes outlined in the plantation establishment model of Burdett (Can J For Res 20:415–427, 1990), but incorporates other physiological processes that drive seedling survival, such as N remobilization, carbohydrate storage and plant hydraulics. The model considers that seedling survival in Mediterranean climates is linked to high growth capacity during the wet season. The model is for container plants and is based on three main principles, (1) Mediterranean climates are not dry the entire year but usually have two seasons of contrasting water availability; (2) summer drought is the main cause of seedling mortality; in this context, deep and large roots is a key trait for avoiding lethal water stress; (3) attainment of large root systems in the dry season is promoted when seedlings have high growth during the wet season. High growth is achieved when seedlings can divert large amount of resources to support new root and shoot growth. Functional traits that confer high photosynthesis, nutrient remobilization capacity, and non-structural carbohydrate storage promote high growth. Increases in seedling size and nutrient concentration strongly affect these physiological processes. Traits that confer high drought resistance are of low value during the wet season because hinder growth capacity. We provide specific evidence to support the model and finally we discuss its implications and the factors that may alter the frequent increase in performance with increase in seedling size and tissue nutrient concentration.


Carbohydrates Drought stress Fertilization Forest plantation Nitrogen Nutrients Photosynthesis Plant quality Remobilization Root growth 



We are grateful to the Centro “El Serranillo” (MAGRAMA), which supported most of the experiments that yielded the results shown in the figures and many other studies referenced in bibliography. Bárbara Cuesta and Mercedes Uscola were supported by FPI-MEC and FPU-MEC grants, respectively. Norberto Heredia received a research contract of the Community of Madrid. This study was also supported by the projects AGL2011-24296 ECOLPIN, CGL2010-18312 (MICIIN), and the network REMEDINAL-2 (S2009/AMB/1783) of the Community of Madrid. We acknowledge D.F. Jacobs for reviewing English writing and two anonymous reviewers whose comments helped improving the paper.


  1. Abod SA, Webster AD, Quinlan JD (1991) Carbohydrates and their effects on the growth and establishment of Tilia and Betula: II. The early season movement of carbohydrates between shoots and roots. J Hort Sci 66:345–355Google Scholar
  2. Adams HD, Guardiola-Claramonte M, Barron-Gafford GA, Villegas JC, Breshears DD, Zou CB, Troch PA, Huxman TE (2009) Temperature sensitivity of drought-induced tree mortality portends increased regional die-off under global-change-type drought. PNAS 106:7063–7066PubMedCrossRefGoogle Scholar
  3. Alía R, Alba N, Agúndez D, Iglesias S (2005) Manual para la comercialización y producción de semillas y plantas forestales. Materiales de base y de reproducción. DGB, MadridGoogle Scholar
  4. Alloza JA, Vallejo R (1999) Relación entre las características meteorológicas del año de plantación y los resultados de las repoblaciones. Ecología 13:173–187Google Scholar
  5. Arnott JT, Grossnickle SC, Puttonen P, Mitchel AK, Folk RS (1993) Influence of nursery culture on growth, cold hardiness, and drought resistance of yellow cypress. Can J For Res 23:2537–2547CrossRefGoogle Scholar
  6. Atzmon N, Reuveni O, Riov J (1994) Lateral root formation in pine seedlings. II The role of assimilates. Trees Struct Funct 8:273–277CrossRefGoogle Scholar
  7. Bayley AD, Kietzka JW (1997) Stock quality and field performance of Pinus patula seedlings produced under two nursery growing regimes during seven different nursery production periods. New For 13:341–356CrossRefGoogle Scholar
  8. Bellot J, Ortiz de Urbina JM, Bonet A, Sánchez JR (2002) The effects of treeshelters on the growth of Quercus coccifera L. seedlings in a semiarid environment. Forestry 75:89–106CrossRefGoogle Scholar
  9. Bernier PY, Lamhamedi MS, Simpson DG (1995) Shoot: Root ratio is of limited use in evaluating the quality of container conifer stock. Tree Planter’s Notes 46:102–106Google Scholar
  10. Brissette JC, Chambers JL (1992) Leaf water status and root system water flux of shortleaf pine (Pinus echinata Mill.) seedlings in relation to new root growth after transplanting. Tree Physiol 11:289–303PubMedGoogle Scholar
  11. Brodribb TJ, Cochard H (2009) Hydraulic failure defines the recovery and point of death in water-stressed conifers. Plant Physiol 149:575–584PubMedCrossRefGoogle Scholar
  12. Bucci SJ et al (2006) Nutrient availability constrains the hydraulic architecture and water relations of savannah trees. Plant Cell Environ 29:2153–2167PubMedCrossRefGoogle Scholar
  13. Burdett AN (1990) Physiological processes in plantation establishment and the development of specifications for forest planting stock. Can J For Res 20:415–427CrossRefGoogle Scholar
  14. Canham CD, Kobe RK, Latty EF, Chazdon RL (1999) Interspecific and intraspecific variation in tree seedling survival: effects of allocation to roots versus carbohydrates reserves. Oecologia 121:1–11CrossRefGoogle Scholar
  15. Castro J (2006) Short delay in timing of emergence determines establishment success in Pinus sylvestris across microhabitats. Ann Bot 98:1233–1240PubMedCrossRefGoogle Scholar
  16. Castro J, Zamora R, Hódar JA, Gómez JM, Gómez Aparicio L (2004) Benefits of using shrubs as nurse plants for reforestation in Mediterranean mountains: a 4-year study. Rest Ecol 12:352–358CrossRefGoogle Scholar
  17. Cerasoli S, Maillard P, Scartazza A, Brugnoli E, Chaves MM, Pereira JS (2004) Carbon and nitrogen winter storage and remobilisation during seasonal flush growth in two-year-old cork oak (Quercus suber L.) saplings. Ann For Sci 61:721–729CrossRefGoogle Scholar
  18. Clearwater MJ, Meinzer FC (2001) Relationships between hydraulic architecture and leaf photosynthetic capacity in nitrogen-fertilized Eucalyptus grandis trees. Tree Physiol 21:683–690PubMedCrossRefGoogle Scholar
  19. Corchero de la Torre S, Gozalo-Cano M, Villar-Salvador P, Peñuelas-Rubira JL (2002) Crecimiento radical en campo de Pinus halepensis y Quercus ilex plantados en diferentes momentos. Revista Montes 68:5–11Google Scholar
  20. Cuesta B, Vega J, Villar-Salvador P, Rey-Benayas JM (2010a) Root growth dynamics of Aleppo pine (Pinus halepensis Mill.) seedlings in relation to shoot elongation, plant size and tissue nitrogen concentration. Trees Struct Funct 24:899–908CrossRefGoogle Scholar
  21. Cuesta B, Villar-Salvador P, Puértolas J, Jacobs D, Rey-Benayas JM (2010b) Why do large, nitrogen rich seedlings better resist stressful transplanting conditions? A physiological analysis in two functionally contrasting Mediterranean forest species. For Ecol Manag 260:71–78CrossRefGoogle Scholar
  22. De Luis M, Verdú M, Raventós J (2008) Early to rise makes a plant healthy, wealthy, and wise. Ecology 89:3061–3071CrossRefGoogle Scholar
  23. del Campo A, Navarro Cerrillo RM, Hermoso J, Ibáñez AJ (2007) Relationships between site and stock quality in Pinus halepensis Mill. reforestation on semiarid landscapes in eastern Spain. Ann For Sci 64:719–731CrossRefGoogle Scholar
  24. del Campo AD, Navarro RM, Ceacero CJ (2010) Seedling quality and field performance of commercial stocklots of containerized holm oak (Quercus ilex) in Mediterranean Spain: an approach for establishing a quality standard. New For 39:17–37CrossRefGoogle Scholar
  25. Dyckmans J, Flessa H (2001) Influence of tree internal N status on uptake and translocation of C and N in beech: a dual 13C and 15 N labeling approach. Tree Physiol 21:395–401PubMedCrossRefGoogle Scholar
  26. El Omari B, Aranda X, Verdaguer D, Pascual G, Fleck I (2003) Resource remobilization in Quercus ilex resprouts. Plant Soil 252:349–357CrossRefGoogle Scholar
  27. Field C, Mooney HA (1986) The photosynthesis-nitrogen relationship in wild plants. In: Givnish TJ (ed) On the economy of plant form and function. Cambridge Univ. Press, Cambridge, pp 25–55Google Scholar
  28. Grelet GA, Alexander IJ, Millard P, Proe MF (2003) Does morphology or the size of the internal nitrogen store determine how Vaccinium spp. respond to spring nitrogen supply? Funct Ecol 17:690–699CrossRefGoogle Scholar
  29. Grossnickle S (2005) Importance of root growth in overcoming planting stress. New For 30:273–294CrossRefGoogle Scholar
  30. Grossnickle S (2012) Why seedlings survive. Influence of plant attributes. New For (this IUFRO issue)Google Scholar
  31. Grossnickle S, Russell JH (1990) Water movement in Yellow-cedar seedlings and rooted cuttings: comparison of whole plant and root system pressurization methods. Tree Physiol 6:57–68PubMedGoogle Scholar
  32. Guehl JM, Falconnet G, Gruez J (1989) Caractéristiques physiologiques et survie après plantation de plants de Cedrus atlantica élevés en conteneurs sur différents types de substrats de culture. Ann Sci For 46:1–14CrossRefGoogle Scholar
  33. Hacke UG, Plavcová L, Almeida-Rodriguez A, King-Jones S, Zhou W, Cooke JEK (2010) Influence of nitrogen fertilization on xylem traits and aquaporin expression in stems of hybrid poplar. Tree Physiol 30:1016–1025PubMedCrossRefGoogle Scholar
  34. Hansen J, Vogg G, Beck E (1996) Assimilation, allocation and utilization of carbon by 3-year-old Scots pine (Pinus sylvestris L.) trees during winter and early spring. Trees Struct Funct 11:83–90Google Scholar
  35. Hernández EI, Vilagrosa A, Luis VC, Llorca M, Chirino E, Vallejo VR (2009) Root hydraulic conductance, gas exchange and leaf water potential in seedlings of Pistacia lentiscus L. and Quercus suber L. grown under different fertilization and light regimes. Environ Exp Bot 67:269–276CrossRefGoogle Scholar
  36. Krasowski MJ, Owens JN (1999) Tracheids in white spruce seedling’s long lateral roots in response to nitrogen availability. Plant Soil 217:215–228CrossRefGoogle Scholar
  37. Leiva MJ, Fernández-Alés R (1998) Variability in seedling water status during drought within a Quercus ilex subsp. ballota population, and its relation to seedling morphology. For Ecol Manag 111:147–156CrossRefGoogle Scholar
  38. Leshem B (1965) The annual activity of intermediary roots of the Aleppo pine. For Sci 11:291–298Google Scholar
  39. Levitt J (1980) Responses of plants to environmental stresses. Volume II. Water, radiation, salt, and other stresses, 2nd edn. Academic Press, New YorkGoogle Scholar
  40. Löf M, Dey DC, Navarro RM, Jacobs DF (2012) Mechanical site preparation for forest restoration. New For (this IUFRO issue)Google Scholar
  41. López B, Sabaté S, Gracia CA (2001) Annual and seasonal changes in fine root biomass of a Quercus ilex L. forest. Plant Soil 230:125–134CrossRefGoogle Scholar
  42. Lovelock CE, Feller IC, McKee KL, Engelbrecht BMJ, Ball MC (2004) The effect of nutrient enrichment on growth, photosynthesis and hydraulic conductance of dwarf mangroves in Panamá. Funct Ecol 18:25–33CrossRefGoogle Scholar
  43. Luis VC, Puértolas J, Climent J, Peters J, González-Rodríguez A, Morales D, Jiménez M (2009) Nursery fertilization enhances survival and physiological status in Canary Island pine (Pinus canariensis) seedlings planted in a semiarid environment. Eur J For Res 128:221–229CrossRefGoogle Scholar
  44. Luis V, Llorca M, Chirino E, Hernández E, Vilagrosa A (2010) Differences in morphology, gas exchange and root hydraulic conductance before planting in Pinus canariensis seedlings growing under different fertilization and light regimes. Trees Struct Funct 24:1143–1150CrossRefGoogle Scholar
  45. Maillard P, Garriou D, Deléens E, Gross P, Guehl JM (2004) The effects of lifting on mobilisation and new assimilation of C and N during regrowth of transplanted Corsican pine seedlings. A dual 13C and 15 N labelling approach. Ann For Sci 61:795–805CrossRefGoogle Scholar
  46. Martínez-Sanz A (2006) Estudio del efecto del volumen del contenedor y la fertilización en vivero sobre la calidad de planta y el establecimiento de los brinzales de Juniperus thurifera L. BSc Thesis. Universidad Politécnica de Madrid, Madrid, p 90Google Scholar
  47. McDowell N, Pockman WT, Allen CD, Breshears DD, Cobb N, Kolb TE, Plaut J, Sperry JS, West A, Williams DG, Yepez EA (2008) Mechanisms of plant survival and mortality during drought: why do some plants survive while others succumb to drought? New Phytol 178:719–739PubMedCrossRefGoogle Scholar
  48. McPherson K, Williams K (1998) The role of carbohydrate reserves in the growth, resilience, and persistence of cabbage palm seedlings (Sabal palmetto). Oecologia 117:460–468CrossRefGoogle Scholar
  49. Mediavilla S, Escudero A (2004) Stomatal responses to drought of mature trees and seedlings of two co-occurring Mediterranean oaks. For Ecol Manag 187:281–294CrossRefGoogle Scholar
  50. Mendoza I, Zamora R, Castro J (2009) A seeding experiment for testing tree-community recruitment under variable environments: implications for forest regeneration and conservation in Mediterranean habitats. Biol Conserv 142:1491–1499CrossRefGoogle Scholar
  51. Mexal JG, Landis TD (1990) Target seedling concepts: height and diameter. In: Rose R, Campbell SJ, Landis TD (eds) Target seedling symposium. UDSA Forest Service, Roseburg (Oregon), pp 17–35Google Scholar
  52. Milla R, Castro-Díez P, Maestro-Martínez M, Montserrat-Martí G (2005) Relationships between phenology and the remobilization of nitrogen, phosphorus and potassium in branches of eight Mediterranean evergreens. New Phytol 168:167–178PubMedCrossRefGoogle Scholar
  53. Millard P, Grelet G (2010) Nitrogen storage and remobilization by trees: ecophysiological relevance in a changing world. Tree Physiol 30:1083–1095PubMedCrossRefGoogle Scholar
  54. Millard P, Neilsen GH (1989) The influence of nitrogen supply on the uptake and remobilisation of stored N for the seasonal growth of apple trees. Ann Bot 63:301–309Google Scholar
  55. Millet J, Millard P, Hester AJ, McDonald AJS (2005) Do competition and herbivory alter the internal nitrogen dynamics of birch saplings? New Phytol 168:413–422CrossRefGoogle Scholar
  56. Mitrakos K (1980) A theory for Mediterranean plant life. Acta Oecol Oecol Plant 1:245–252Google Scholar
  57. Navarro RM, Villar-Salvador P, del Campo A (2006) Morfología y establecimiento de los plantones. In: Cortina J, Peñuelas JL, Puértolas J, Savé R, Vilagrosa A (eds) Calidad de planta forestal para la restauración en ambientes mediterráneos degradados. Estado actual de conocimientos. Ministerio de Medio Ambiente, Madrid, pp 67–88Google Scholar
  58. Oliet J, Planelles R, Artero F, Valverde R, Jacobs D, Segura M (2009) Field performance of Pinus halepensis planted in Mediterranean arid conditions: relative influence of seedling morphology and mineral nutrition. New For 37:313–331CrossRefGoogle Scholar
  59. Padilla FM, Pugnaire FI (2007) Rooting depth and soil moisture control Mediterranean woody seedling survival during drought. Funct Ecol 21:489–495CrossRefGoogle Scholar
  60. Palacios G, Navarro Cerrillo RM, del Campo A, Toral M (2009) Site preparation, stock quality and planting date effect on early establishment of Holm oak (Quercus ilex L.) seedlings. Ecol Eng 35:38–46CrossRefGoogle Scholar
  61. Pinto JR, Marshall JD, Dumroese RK, Davis AS, Cobos DR (2012) Photosynthetic response, carbon isotopic composition, survival, and growth of three stock types under water stress enhanced by vegetative competition. Can J For Res 42:333–344CrossRefGoogle Scholar
  62. Planelles González R (2004) Efectos de la fertilización NPK en vivero sobre la calidad funcional de plantas de Ceratonia siliqua L. PhD Sci Thesis. Universidad Politécnica de Madrid, MadridGoogle Scholar
  63. Puértolas J, Gil L, Pardos JA (2003) Effects of nutritional status and seedling size on fiel performance of Pinus halepensis planted on former arable land in th Mediterranean basin. Forestry 76:159–168CrossRefGoogle Scholar
  64. Puttonen P (1986) Carbohydrate reserves in Pinus sylvestris seedling needles as an attribute of seedling vigor. Scand J For Res 1:181–193Google Scholar
  65. Ramírez-Valiente JA, Valladares F, Gil L, Aranda I (2009) Population differences in juvenile survival under increasing drought are mediated by seed size in cork oak (Quercus suber L.). For Ecol Manag 257:1676–1683CrossRefGoogle Scholar
  66. Rodríguez-García B (2003) Estudio del momento óptimo de plantación de brinzales de Quercus ilex y Pinus halepensis en función del clima de la estación de plantación. BSc Thesis. Universidad Politécnica de Madrid, MadridGoogle Scholar
  67. Sala A, Tenhunen JD (1994) Site-specific water relations and stomatal response of Quercus ilex in a Mediterranean watershed. Tree Physiol 14:601–617PubMedGoogle Scholar
  68. Salifu KF, Timmer VR (2003) Nitrogen retranslocation response of young Picea mariana to nitrogen-15 supply. Soil Sci Soc Am J 67:309–317CrossRefGoogle Scholar
  69. Silla F, Escudero A (2003) Uptake, demand and internal cycling of nitrogen in saplings of Mediterranean Quercus species. Oecologia 136:28–36PubMedCrossRefGoogle Scholar
  70. Sloan JL, Jacobs D (2008) Carbon translocation patterns associated with new root proliferation during episodic growth of transplanted Quercus rubra seedlings. Tree Physiol 28:1121–1126PubMedCrossRefGoogle Scholar
  71. South DB, Larsen HS, Williams HM, Boyer JN (1989) Use of seedling size as a covariate for root growth potential studies. In: 5th Biennial southern Silvicultural research conference, pp 89–93Google Scholar
  72. South DB, Harris SW, Barnett JP, Hainds MJ, Gjerstad DH (2005) Effect of container type and seedling size on survival and early height growth of Pinus palustris seedlings in Alabama, USA. For Ecol Manag 204:385–398CrossRefGoogle Scholar
  73. Sperry JS (2000) Hydraulic constraints on plant gas exchange. Agric For Meteorol 104:13–23CrossRefGoogle Scholar
  74. Sword Sayer MA, Brissette JC, Barnett JP (2005) Root growth and hydraulic conductivity of southern pine seedlings in response to soil temperature and water availability after planting. New For 30:253–272CrossRefGoogle Scholar
  75. Tan W, Hogan GD (1997) Physiological and morphological responses to nitrogen limitation in jack pine seedlings: potential implications for drought tolerance. New For 14:19–31CrossRefGoogle Scholar
  76. Thompson BE (1985) Seedling morphological evaluation. What you can tell by looking. In: Duryea ML (ed) Evaluating seedling quality: principles, procedures, and predictive abilities of major tests. Oregon State University, Corvallis (Oregon), pp 59–71Google Scholar
  77. Trubat R, Cortina J, Vilagrosa A (2006) Plant morphology and root hydraulics are altered by nutrient deficiency in Pistacia lentiscus L. Trees Struct Funct 20:334–339CrossRefGoogle Scholar
  78. Trubat R, Cortina J, Vilagrosa A (2011) Nutrient deprivation improves field performance of woody seedlings in a degraded semi-arid shrubland. Ecol Eng 37:1164–1173CrossRefGoogle Scholar
  79. Tsakaldimi M, Zagas T, Tsitsoni T, Ganatsas P (2005) Root morphology, stem growth and field performance of seedlings of two Mediterranean evergreen oak species raised in different container types. Plant Soil 278:85–93CrossRefGoogle Scholar
  80. Tuttle CL, South DB, Golden MS, Meldahl RS (1988) Initial Pinus taeda seedling height relationships with early survival and growth. Can J For Res 18:867–871CrossRefGoogle Scholar
  81. van den Driessche R (1987) Importance of current photosynthate to new root growth in planted conifer seedlings. Can J For Res 17:776–782CrossRefGoogle Scholar
  82. van den Driessche R (1991a) Influence of container nursery regimes on drought resistance of seedlings following planting. I. Survival and growth. Can J For Res 21:555–565CrossRefGoogle Scholar
  83. van den Driessche R (1991b) New root growth of Douglas-fir seedlings at low carbon dioxide concentration. Tree Physiol 8:289–295Google Scholar
  84. van den Driessche R (1992) Changes in drought resistance and root growth capacity of container seedlings in response to nursery drought, nitrogen, and potassium treatments. Can J For Res 22:740–749CrossRefGoogle Scholar
  85. Verdaguer D, Vilagran J, Lloansi S, Fleck I (2011) Morphological and physiological acclimation of Quercus coccifera L. seedlings to water availability and growing medium. New For 42:363–381CrossRefGoogle Scholar
  86. Vilagrosa A, Bellot J, Vallejo VR, Gil Pelegrín E (2003) Cavitation, stomatal conductance, and leaf dieback in seedlings of two co-occurring Mediterranean shrubs during an intense drought. J Exp Bot 54:2015–2024PubMedCrossRefGoogle Scholar
  87. Villar-Salvador P (2003) Importancia de la calidad de las plantas en los proyectos de revegetación. In: Rey Benayas JM, Espigares, T, y Nicolau JM (eds) Restauración de ecosistemas mediterráneos. Publicaciones de la Universidad de Alcalá, Alcalá de Henares, pp 65–86Google Scholar
  88. Villar-Salvador P, Planelles R, Enríquez E, Peñuelas Rubira J (2004a) Nursery cultivation regimes, plant functional attributes, and field performance relationships in the Mediterranean oak Quercus ilex L. For Ecol Manag 196:257–266CrossRefGoogle Scholar
  89. Villar-Salvador P, Planelles R, Oliet J, Peñuelas-Rubira JL, Jacobs DF, González M (2004b) Drought tolerance and transplanting performance of holm oak (Quercus ilex) seedlings after drought hardening in the nursery. Tree Physiol 24:1147–1155PubMedCrossRefGoogle Scholar
  90. Villar-Salvador P, Puértolas J, Planelles R, Peñuelas Rubira J (2005) Effect of nitrogen fertilization in the nursery on the drought and frost resistance of Mediterranean forest species. For Syst (formerly Inv Agrarias: Sist Rec For) 14:408–418Google Scholar
  91. Willaume M, Pages L (2006) How periodic growth pattern and source/sink relations affect root growth in oak tree seedlings. J Exp Bot 57:815–826PubMedCrossRefGoogle Scholar
  92. Zou C, Penfold C, Sands R, Misra RK, Hudson I (2001) Effects of soil air-filled porosity, soil matric potential and soil strength on primary root growth of radiata pine seedlings. Plant Soil 236:105–115CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2012

Authors and Affiliations

  • Pedro Villar-Salvador
    • 1
    Email author
  • Jaime Puértolas
    • 2
    • 3
  • Bárbara Cuesta
    • 1
  • Juan L. Peñuelas
    • 2
  • Mercedes Uscola
    • 1
  • Norberto Heredia-Guerrero
    • 1
  • José M. Rey Benayas
    • 1
  1. 1.Departamento de EcologíaUniversidad de AlcaláMadridSpain
  2. 2.Centro Nacional de Recursos Genéticos Forestales “El Serranillo”Ministerio de AgriculturaGuadalajaraSpain
  3. 3.Lancaster Environment CentreLancaster UniversityLancasterUK

Personalised recommendations