Drought impact on carbon and water cycling in a Mediterranean Quercus suber L. woodland during the extreme drought event in 2012

Biogeosciences

Volumn 11, Issue 24, 2014, Pages 7159-7178

  Piayda, A ^a   Dubbert, M ^b   Rebmann, C ^a   Kolle, O ^c   Costa E Silva, F ^d   Correia, A ^d   Pereira, J S ^d   Werner, C ^b   Cuntz, M ^a  

^a HELMHOLTZ CENTRE FOR ENVIRONMENTAL RESEARCH UFZ   (Germany)

^b UNIVERSITY OF BAYREUTH   (Germany)

^c MAX PLANCK INSTITUTE FOR BIOGEOCHEMISTRY   (Germany)

^d INSTITUTO SUPERIOR DE AGRONOMIA   (Portugal)

Author keywords

[No Author keywords available]

Indexed keywords

CARBON SINK; DECIDUOUS TREE; DROUGHT STRESS; ECOLOGICAL MODELING; HYDROLOGICAL CYCLE; MORTALITY; PHOTOSYNTHESIS; PRECIPITATION (CLIMATOLOGY); PRIMARY PRODUCTION; WOODLAND;

QUERCUS SUBER;

EID: 84944192438     PISSN: 17264170     EISSN: 17264189     Source Type: Journal    
DOI: 10.5194/bg-11-7159-2014     Document Type: Article

References (119)

1. Aubinet, M., Grelle, A., Ibrom, A., Rannik, Ü., Moncrieff, J., Foken, T., Kowalski, A., Martin, P., Berbigier, P., Bernhofer, C., Clement, R., Elbers, J., Granier, A., Grünwald, T., Morgenstern, K., Pilegaard, K., Rebmann, C., Snijders, W., Valentini, R., and Vesala, T.: Estimates of the Annual Net Carbon and Water Exchange of Forests: The EUROFLUX Methodology, vol. 30 of Adv. Ecol. Res., Academic Press, 113-175, doi:10.1016/S0065-2504(08)60018-5, 1999.
2. Baldocchi, D. D., Ma, S., Rambal, S., Misson, L., Ourcival, J.-M., Limousin, J.-M., Pereira, J., and Papale, D.: On the differential advantages of evergreenness and deciduousness in mediterranean oak woodlands: a flux perspective, Ecol. Appl., 20, 1583-1597, doi:10.1890/08-2047.1, 2009.
3. Ball, J., Woodrow, L. E., and Beny, J. A.: A model predicting stomatal conductance and its contribution to the control of photosynthesis under different environmental conditions, in: Progress in Photosynthesis Research, edited by: Biggins, J., vol. 4, Nijhoff, Dordrecht, 221-224, 1987.
4. Beer, C., Reichstein, M., Tomelleri, E., Ciais, P., Jung, M., Carvalhais, N., Rödenbeck, C., Arain, M. A., Baldocchi, D., Bonan, G. B., Bondeau, A., Cescatti, A., Lasslop, G., Lindroth, A., Lomas, M., Luyssaert, S., Margolis, H., Oleson, K. W., Roupsard, O., Veenendaal, E., Viovy, N., Williams, C., Woodward, F. I., and Papale, D.: Terrestrial gross carbon dioxide uptake: global distribution and covariation with climate, Science, 329, 834-838, doi:10.1126/science.1184984, 2010.
5. 3plants for use in dynamic global vegetation models, Glob. Change Biol., 1, 289-294, doi:10.1111/j.1365-2486.1995.tb00027.x, 1995.
6. Berry, J. and Björkman, O.: Photosynthetic response and adaptation to temperature in higher plants, Ann. Rev. Plant Physio., 31, 491-543, doi:10.1146/annurev.pp.31.060180.002423, 1980.
7. Besson, C. K., do Vale, R. L., Rodrigues, M. L., Almeida, P., Herd, A., Grant, O. M., David, T. S., Schmidt, M., Otieno, D., Keenan, T. F., Gouveia, C., Mériaux, C., Chaves, M. M., and Pereira, J. S.: Cork oak physiological responses to manipulated water availability in a Mediterranean woodland, Agr. Forest Meteorol., 184, 230-242, doi:10.1016/j.agrformet.2013.10.004, 2014.
8. Beyschlag, W., Lange, O., and Tenhunen, J. D.: Photosynthesis and water relations of the mediterranean evergreen sclerophyll Arbutus unedo L. throughout the year at a site in Portugal. I. diurnal courses of carbon dioxide gas exchange and transpiration under natural conditions, Flora, 178, 409-444, 1986.
9. Bissinger, V. and Kolditz, O.: Helmholtz Interdisciplinary Graduate School for Environmental Research (HIGRADE), GAIA, 1, 71-73, 2008.
10. Bonan, G. B.: Ecological Climatology: Concepts and Applications, Cambridge University Press, 2002.
11. Bugalho, M. N., Caldeira, M. C., Pereira, J. S., Aronson, J., and Pausas, J. G.: Mediterranean cork oak savannas require human use to sustain biodiversity and ecosystem services, Front. Ecol. Environ., 9, 278-286, doi:10.1890/100084, 2011.
12. Bussotti, F., Ferrini, F., Pollastrini, M., and Fini, A.: The challenge of Mediterranean sclerophyllous vegetation under climate change: from acclimation to adaptation, Environ. Exp. Bot., 103, 80-98, doi:10.1016/j.envexpbot.2013.09.013, 2013.
13. Chaves, M. M., Pereira, J. S., Maroco, J., Rodrigues, M. L., Ricardo, C. P. P., Osório, M. L., Catvalho, I., Faria, T., and Pinheiro, C.: How plants cope with water stress in the field? Photosynthesis and growth, Ann. Bot.-London, 89, 907-916, doi:10.1093/aob/mcf105, 2002.
14. Ciais, P., Reichstein, M., Viovy, N., Granier, A., Ogee, J., Allard, V., Aubinet, M., Buchmann, N., Bernhofer, C., Carrara, A., Chevallier, F., De Noblet, N., Friend, A. D., Friedlingstein, P., Grunwald, T., Heinesch, B., Keronen, P., Knohl, A., Krinner, G., Loustau, D., Manca, G., Matteucci, G., Miglietta, F., Ourcival, J. M., Papale, D., Pilegaard, K., Rambal, S., Seufert, G., Soussana, J. F., Sanz, M. J., Schulze, E. D., Vesala, T., and Valentini, R.: Europe-wide reduction in primary productivity caused by the heat and drought in 2003, Nature, 437, 529-533, doi:10.1038/nature03972, 2005.
15. 4plants, Aust. J. Plant Physiol., 19, 519-538, doi:10.1071/PP9920519, 1992.
16. Costa, A. C., Santos, J. A., and Pinto, J. G.: Climate change scenarios for precipitation extremes in Portugal, Theor. Appl. Climatol., 108, 217-234, doi:10.1007/s00704-011-0528-3, 2012.
17. Costa e Silva, F., Correia, A. C., Correia, A. V., Piayda, A., Dubbert, M., Werner, C., David, J. S., and Pereira, J. S.: Quercus suber phenological adjustments to seasonal water availability: influence on net ecosystem exchange, Agr. Forest Meteorol., submitted, 2014.
18. Cowan, I.: Stomatal Behaviour and Environment, Academic Press, available at: http://books.google.com.au/books?id=GYhvNQAACAAJ (last access: 1 July 2014), 1977.
19. Damour, G., Simonneau, T., Cochard, H., and Urban, L.: An overview of models of stomatal conductance at the leaf level, Plant Cell Environ., 33, 1419-1438, doi:10.1111/j.1365-3040.2010.02181.x, 2010.
20. David, T., Ferreira, M., Cohen, S., Pereira, J., and David, J.: Constraints on transpiration from an evergreen oak tree in southern Portugal, Agr. Forest Meteorol., 122, 193-205, doi:10.1016/j.agrformet.2003.09.014, 2004.
21. David, T. S., Henriques, M. O., Kurz-Besson, C., Nunes, J., Valente, F., Vaz, M., Pereira, J. S., Siegwolf, R., Chaves, M. M., Gazarini, L. C., and David, J. S.: Water-use strategies in two co-occurring Mediterranean evergreen oaks: surviving the summer drought, Tree Physiol., 27, 793-803, doi:10.1093/treephys/27.6.793, 2007.
22. de Dios Miranda, J., Padilla, F. M., and Pugnaire, F. I.: Response of a Mediterranean semiarid community to changing patterns of water supply, Perspect. Plant Ecol., 11, 255-266, doi:10.1016/j.ppees.2009.05.001, 2009.
23. Demmig-Adams, B. and Adams, W. W.: Photoprotection and other responses of plants to high light stress, Ann. Rev. Plant Biol., 43, 599-626, 1992.
24. De Pury, D. G. G. and Farquhar, G. D.: Simple scaling of photosynthesis from leaves to canopies without the errors of big-leaf models, Plant Cell Environ., 20, 537-557, doi:10.1111/j.1365-3040.1997.00094.x, 1997.
25. Dubbert, M., Mosena, A., Piayda, A., Cuntz, M., Correia, A., Pereira, J. S., and Werner, C.: Influence of tree cover on herbaceous layer development and fluxes in a Portuguese cork oak woodland, Acta Oecol., 59, 35-45, 2014a.
26. Dubbert, M., Piayda, A., Cuntz, M., Correia, A. C., Costa e Silva, F., Pereira, J. S., andWerner, C.: Stable oxygen isotope and flux partitioning d emonstrates understory of an oak savanna contributes up to half of ecosystem carbon and water exchange, Frontiers in Plant Science, 5, 1-16, doi:10.3389/fpls.2014.00530, 2014b.
27. Duckstein, L.: Multiobjective optimization in structural design: the model choice problem, in: New Directions in Optimum Structural Design, edited by: Atrek, E., John Wiley, 1981.
28. Egea, G., Verhoef, A., and Vidale, P. L.: Towards an improved and more flexible representation of water stress in coupled photosynthesis-stomatal conductance models, Agr. Forest Meteorol., 151, 1370-1384, doi:10.1016/j.agrformet.2011.05.019, 2011.
29. Ehleringer, J. R. and Cook, C. S.: Photosynthesis in encelia farinosa gray in response to decreasing leaf water potential, Plant Physiol., 75, 688-693, doi:10.1104/pp.75.3.688, 1984.
30. Espigares, T. and Peco, B.: Mediterranean pasture dynamics: the role of germination, J. Veg. Sci., 4, 189-194, 1993.
31. Espigares, T. and Peco, B.: Mediterranean annual pasture dynamics: impact of autumn drought, J. Ecol., 83, 135-142, 1995.
32. 2flux, Bound.-Lay. Meteorol., 74, 321-340, doi:10.1007/BF00712375, 1995.
33. 3species, Planta, 149, 78-90, doi:10.1007/BF00386231, 1980.
34. Farquhar, G. D. and Sharkey, T. D.: Stomatal conductance and photosynthesis, Ann. Rev. Plant Physio., 33, 317-345, doi:10.1146/annurev.pp.33.060182.001533, 1982.
35. Field, C., Barros, V., Stocker, T., Qin, D., Dokken, D., Ebi, K., Mastrandrea, M., Mach, K., Plattner, G.-K., Allen, S., Tignor, M., and Midgley, P. (Eds.): Managing the Risks of Extreme Events and Disasters to Advance Climate Change Adaptation, IPCC Special Reports, Cambridge University Press, Cambridge, available at: https://www.ipcc.ch/pdf/special-reports/srex/SREX-Full-Report.pdf, 2012.
36. Figueroa, M. E. and Davy, A. J.: Response of Mediterranean grass-land species to changing rainfall, J. Ecol., 79, 925-941, 1991.
37. Fischer, G., van Velthuizen, H., Shah, M., and Nachtergaele, F. (Eds.): Global Agro-ecological Assessment for Agriculture in the 21st Century: Methodology and Results, International Institute for Applied Systems Analysis, 2002.
38. Foken, T.: The energy balance closure problem: an overview, Ecol. Appl., 18, 1351-1367, doi:10.1890/06-0922.1, 2008.
39. Foken, T. and Wichura, B.: Tools for quality assessment of surface-based flux measurements, Agr. Forest Meteorol., 78, 83-105, doi:10.1016/0168-1923(95)02248-1, 1996.
40. García-Barrón, L., Morales, J., and Sousa, A.: Characterisation of the intra-annual rainfall and its evolution (1837-2010) in the southwest of the Iberian Peninsula, Theor. Appl. Climatol., 114, 445-457, doi:10.1007/s00704-013-0855-7, 2013.
41. Gordo, O. and Sanz, J. J.: Phenology and climate change: a long-term study in a Mediterranean locality, Oecologia, 146, 484-495, doi:10.1007/s00442-005-0240-z, 2005.
42. Grace, J., José, J. S., Meir, P., Miranda, H. S., and Montes, R. A.: Productivity and carbon fluxes of tropical savannas, J. Biogeogr., 33, 387-400, doi:10.1111/j.1365-2699.2005.01448.x, 2006.
43. Granier, A., Reichstein, M., Bréda, N., Janssens, I., Falge, E., Ciais, P., Grünwald, T., Aubinet, M., Berbigier, P., Bernhofer, C., Buchmann, N., Facini, O., Grassi, G., Heinesch, B., Ilvesniemi, H., Keronen, P., Knohl, A., Köstner, B., Lagergren, F., Lindroth, A., Longdoz, B., Loustau, D., Mateus, J., Montagnani, L., Nys, C., Moors, E., Papale, D., Peiffer, M., Pilegaard, K., Pita, G., Pumpanen, J., Rambal, S., Rebmann, C., Rodrigues, A., Seufert, G., Tenhunen, J., Vesala, T., and Wang, Q.: Evidence for soil water control on carbon and water dynamics in European forests during the extremely dry year: 2003, Agr. Forest Meteorol., 143, 123-145, doi:10.1016/j.agrformet.2006.12.004, 2007.
44. Grant, O. M., Tronina, Ł., Ramalho, J. C., Kurz Besson, C., Lobo-do-Vale, R., Santos Pereira, J., Jones, H. G., and Chaves, M. M.: The impact of drought on leaf physiology of Quercus suber L. trees: comparison of an extreme drought event with chronic rainfall reduction, J. Exp. Bot., 61, 4361-4371, doi:10.1093/jxb/erq239, 2010.
45. Guerreiro, S. B., Kilsby, C. G., and Serinaldi, F.: Analysis of time variation of rainfall in transnational basins in Iberia: abrupt changes or trends?, Int. J. Climatol., 34, 114-133, doi:10.1002/joc.3669, 2013.
46. Heimann, M. and Reichstein, M.: Terrestrial ecosystem carbon dynamics and climate feedbacks, Nature, 451, 289-292, doi:10.1038/nature06591, 2008.
47. Hollinger, D. Y., Kelliher, F. M., Byers, J. N., Hunt, J. E., McSeveny, T. M., and Weir, P. L.: Carbon dioxide exchange between an undisturbed old-growth temperate forest and the atmosphere, Ecology, 75, 134-150, 1994.
48. Hulme, M., Mitchell, J., Ingram, W., Lowe, J., Johns, T., New, M., and Viner, D.: Climate change scenarios for global impacts studies, Global Environ. Chang., 9, S3-S19, doi:10.1016/S0959-3780(99)00015-1, 1999.
49. Huxman, T. E., Wilcox, B. P., Breshears, D. D., Scott, R. L., Snyder, K. A., Small, E. E., Hultine, K., Pockman, W. T., and Jackson, R. B.: Ecohydrological implications of woody plant encroachment, Ecology, 86, 308-319, doi:10.1890/03-0583, 2005.
50. Ibrom, A., Dellwik, E., Larsen, S. E., and Pilegaard, K.: On the use of the Webb-Pearman-Leuning theory for closed-path eddy correlation measurements, Tellus B, 59, 937-946, doi:10.1111/j.1600-0889.2007.00311.x, 2007.
51. Johnson, F. H., Eyring, H., and Williams, R. W.: The nature of enzyme inhibitions in bacterial luminescence: Sulfanilamide, urethane, temperature and pressure, J. Cell. Compar. Physl., 20, 247-268, doi:10.1002/jcp.1030200302, 1942.
52. Jongen, M., Pereira, J. S., Aires, L. M. I., and Pio, C. A.: The effects of drought and timing of precipitation on the inter-annual variation in ecosystem-atmosphere exchange in a Mediterranean grassland, Agr. Forest Meteorol., 151, 595-606, doi:10.1016/j.agrformet.2011.01.008, 2011.
53. Jongen, M., Lecomte, X., Unger, S., Fangueiro, D., and Pereira, J. S.: Precipitation variability does not affect soil respiration and nitrogen dynamics in the understorey of a Mediterranean oak woodland, Plant Soil, 372, 235-251, doi:10.1007/s11104-013-1728-7, 2013a.
54. Jongen, M., Lecomte, X., Unger, S., Pintó-Marijuan, M., and Pereira, J. S.: The impact of changes in the timing of precipitation on the herbaceous understorey of Mediterranean evergreen oak woodlands, Agr. Forest Meteorol., 171-172, 163-173, doi:10.1016/j.agrformet.2012.11.020, 2013b.
55. Jongen, M., Unger, S., Fangueiro, D., Cerasoli, S., Silva, J., and Pereira, J. S.: Resilience of montado understorey to experimental precipitation variability fails under severe natural drought, Agr. Ecosyst. Environ., 178, 18-30, doi:10.1016/j.agee.2013.06.014, 2013c.
56. June, T., Evans, J. R., and Farquhar, G. D.: A simple new equation for the reversible temperature dependence of photosynthetic electron transport: a study on soybean leaf, Funct. Plant Biol., 31, 275-283, doi:10.1071/FP03250, 2004.
57. Kattge, J. and Knorr, W.: Temperature acclimation in a biochemical model of photosynthesis: a reanalysis of data from 36 species, Plant Cell Environ., 30, 1176-1190, doi:10.1111/j.1365-3040.2007.01690.x, 2007.
58. 2exchanges of the terrestrial biosphere: process-based simulations and uncertainties, Global Ecol. Biogeogr., 9, 225-252, doi:10.1046/j.1365-2699.2000.00159.x, 2000.
59. Kolle, O. and Rebmann, C.: EddySoft Documentation of a Software Package to Acquire and Process Eddy Covariance Data, Technical Reports 10, Max-Planck-Institut für Biogeochemie, Jena, available at: http://www.bgc-jena.mpg.de/bgc-processes/staff/corinna.rebmann/tech-report10.pdf, 2007.
60. Krinner, G., Viovy, N., de Noblet-Ducoudré, N., Ogée, J., Polcher, J., Friedlingstein, P., Ciais, P., Sitch, S., and Prentice, I. C.: A dynamic global vegetation model for studies of the coupled atmosphere-biosphere system, Global Biogeochem. Cy., 19, GB1015, doi:10.1029/2003GB002199, 2005.
61. Krishnan, P., Meyers, T. P., Scott, R. L., Kennedy, L., and Heuer, M.: Energy exchange and evapotranspiration over two temperate semi-arid grasslands in North America, Agr. Forest Meteorol., 153, 31-44, doi:10.1016/j.agrformet.2011.09.017, 2012.
62. Kurz-Besson, C., Otieno, D., Lobo do Vale, R., Siegwolf, R., Schmidt, M., Herd, A., Nogueira, C., David, T., David, J., Tenhunen, J., Pereira, J., and Chaves, M.: Hydraulic lift in cork oak trees in a savannah-type Mediterranean ecosystem and its contribution to the local water balance, Plant Soil, 282, 361-378, doi:10.1007/s11104-006-0005-4, 2006.
63. Lasslop, G., Reichstein, M., Papale, D., Richardson, A. D., Arneth, A., Barr, A., Stoy, P., and Wohlfahrt, G.: Separation of net ecosystem exchange into assimilation and respiration using a light response curve approach: critical issues and global evaluation, Glob. Change Biol., 16, 187-208, doi:10.1111/j.1365-2486.2009.02041.x, 2010.
64. 3plants, Plant Cell Environ., 18, 339-355, doi:10.1111/j.1365-3040.1995.tb00370.x, 1995.
65. Leuning, R.: The correct form of the Webb, Pearman and Leuning equation for eddy fluxes of trace gases in steady and non-steady state, horizontally homogeneous flows, Bound.-Lay. Meteorol., 123, 263-267, doi:10.1007/s10546-006-9138-5, 2007.
66. Liu, Y., Zhou, Y., Ju, W., Wang, S., Wu, X., He, M., and Zhu, G.: Impacts of droughts on carbon sequestration by China's terrestrial ecosystems from 2000 to 2011, Biogeosciences, 11, 2583-2599, doi:10.5194/bg-11-2583-2014, 2014.
67. Long, S., Humphries, S., and Falkowski, P. G.: Photoinhibition of photosynthesis in nature, Ann. Rev. Plant Biol., 45, 633-662, 1994.
68. Ma, S., Baldocchi, D. D., Xu, L., and Hehn, T.: Inter-annual variability in carbon dioxide exchange of an oak/grass savanna and open grassland in California, Agr. Forest Meteorol., 147, 157-171, doi:10.1016/j.agrformet.2007.07.008, 2007.
69. Matthews, M. A. and Boyer, J. S.: Acclimation of photosynthesis to low leaf water potentials, Plant Physiol., 74, 161-166, doi:10.1104/pp.74.1.161, 1984.
70. Mauder, M. and Foken, T.: Documentation and Instruction Manual of the Eddy-Covariance Software Package TK3, Universität Bayreuth Abt. Mikrometeorologie, available at: http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:bvb:703-opus-8665, 2011.
71. Mauder, M., Cuntz, M., Drüe, C., Graf, A., Rebmann, C., Schmid, H. P., Schmidt, M., and Steinbrecher, R.: A strategy for quality and uncertainty assessment of long-term eddy-covariance measurements, Agr. Forest Meteorol., 169, 122-135, doi:10.1016/j.agrformet.2012.09.006, 2013.
72. Medlyn, B. E., Dreyer, E., Ellsworth, D., Forstreuter, M., Harley, P. C., Kirschbaum, M. U. F., Le Roux, X., Montpied, P., Strassemeyer, J., Walcroft, A., Wang, K., and Loustau, D.: Temperature response of parameters of a biochemically based model of photosynthesis. II. A review of experimental data, Plant Cell Environ., 25, 1167-1179, doi:10.1046/j.1365-3040.2002.00891.x, 2002.
73. Miranda, P., Coelho, F., Tomé, A. R, Carvalho, A., Pires, C., Pires, H., Pires, V. C., and Ramalho, C.: 20th century Portuguese climate and climate scenarios, in: Climate Change in Portugal: Scenarios, Impacts and Adaptation Measures (SIAM Project), Gradiva, 23-83, 2002.
74. Mourato, S., Moreira, M., and Corte-Real, J.: Interannual variability of precipitation distribution patterns in Southern Portugal, Int. J. Climatol., 30, 1784-1794, doi:10.1002/joc.2021, 2010.
75. Nelder, J. A. and Mead, R.: A simplex method for function minimization, Comput. J., 7, 308-313, doi:10.1093/comjnl/7.4.308, 1965.
76. Oleson, K. W., Lawrence, D. M., Bonan, G. B., Flanner, M. G., Kluzek, E., Lawrence, P. J., Levis, S., Swenson, S. C., and Thornton, P. E.: Technical Description of version 4.0 of the Community Land Model (CLM)leu, Tech. rep., National Center for Atmospheric Research, 2010.
77. Oliveira, G., Correia, O., Martins-Louc¸ão, M., and Catarino, F.: Water relations of cork-oak (Quercus suber L.) under natural conditions, in: Quercus ilex L. Ecosystems: Function, Dynamics and Management, edited by: Romane, F., and Terradas, J., vol. 13 of Advances in vegetation science, Springer Netherlands, 199-208, doi:10.1007/978-94-017-2836-2-21, 1992.
78. Paco, T. A., David, T. S., Henriques, M. O., Pereira, J. S., Valente, F., Banza, J., Pereira, F. L., Pinto, C., and David, J. S.: Evapotranspiration from a Mediterranean evergreen oak savannah: the role of trees and pasture, J. Hydrol., 369, 98-106, doi:10.1016/j.jhydrol.2009.02.011, 2009.
79. Papale, D., Reichstein, M., Aubinet, M., Canfora, E., Bernhofer, C., Kutsch, W., Longdoz, B., Rambal, S., Valentini, R., Vesala, T., and Yakir, D.: Towards a standardized processing of Net Ecosystem Exchange measured with eddy covariance technique: algorithms and uncertainty estimation, Biogeosciences, 3, 571-583, doi:10.5194/bg-3-571-2006, 2006.
80. Paredes, D., Trigo, R. M., Garcia-Herrera, R., and Trigo, I. F.: Understanding precipitation changes in Iberia in early spring: weather typing and storm-tracking approaches, J. Hydrometeorol., 7, 101-113, doi:10.1175/JHM472.1, 2006.
81. Peñuelas, J., Filella, I., and Comas, P.: Changed plant and animal life cycles from 1952 to 2000 in the Mediterranean region, Glob. Change Biol., 8, 531-544, doi:10.1046/j.1365-2486.2002.00489.x, 2002.
82. Peñuelas, J., Filella, I., Zhang, X., Llorens, L., Ogaya, R., Lloret, F., Comas, P., Estiarte, M., and Terradas, J.: Complex spatiotemporal phenological shifts as a response to rainfall changes, New Phytol., 161, 837-846, doi:10.1111/j.1469-8137.2004.01003.x, 2004.
83. Peco, B. and Espigares, T.: Floristic fluctuations in annual pastures: the role of competition at the regeneration stage, J. Veg. Sci., 5, 457-462, doi:10.2307/3235971, 1994.
84. Pereira, J. S., Mateus, J. A., Aires, L. M., Pita, G., Pio, C., David, J. S., Andrade, V., Banza, J., David, T. S., Pac¸o, T. A., and Rodrigues, A.: Net ecosystem carbon exchange in three contrasting Mediterranean ecosystems - the effect of drought, Biogeosciences, 4, 791-802, doi:10.5194/bg-4-791-2007, 2007.
85. Pérez-Ramos, I. M., Rodríguez-Calcerrada, J., Ourcival, J. M., and Rambal, S.: Quercus ilex recruitment in a drier world: a multistage demographic approach, Perspect. Plant Ecol., 15, 106-117, doi:10.1016/j.ppees.2012.12.005, 2013.
86. Piayda, A., Dubbert, M., Werner, C., Correia, A. V., Pereira, J. S., and Cuntz, M.: Quantifying the influence of wooden tissue and leaf clumping on vertically resolved leaf area index and angular gap probability estimates in an open savannah-type woodland, Forest Ecology and Management, submitted 2014.
87. Piñol, J., Lledó, M. J., and Escarré, A.: Hydrological balance of two Mediterranean forested catchments (Prades, northeast Spain), Hydrolog. Sci. J., 36, 95-107, doi:10.1080/02626669109492492, 1991.
88. Rebmann, C., Kolle, O., Heinesch, B., Queck, R., Ibrom, A., and Aubinet, M.: Data acquisition and flux calculations, in: Eddy Covariance: A Practical Guide to Measurement and Data Analysis, Springer, Dordrecht, 59-84, 2012.
89. 2O fluxes at three Mediterranean evergreen sites: revision of current hypotheses?, Glob. Change Biol., 8, 999-1017, doi:10.1046/j.1365-2486.2002.00530.x, 2002.
90. 2O exchange in three Mediterranean ecosystems, J. Geophys. Res.-Atmos., 108, 4726, doi:10.1029/2003JD003430, 2003.
91. Reichstein, M., Falge, E., Baldocchi, D., Papale, D., Aubinet, M., Berbigier, P., Bernhofer, C., Buchmann, N., Gilmanov, T., Granier, A., Grünwald, T., Havránková, K., Ilvesniemi, H., Janous, D., Knohl, A., Laurila, T., Lohila, A., Loustau, D., Matteucci, G., Meyers, T., Miglietta, F., Ourcival, J.-M., Pumpanen, J., Rambal, S., Rotenberg, E., Sanz, M., Tenhunen, J., Seufert, G., Vaccari, F., Vesala, T., Yakir, D., and Valentini, R.: On the separation of net ecosystem exchange into assimilation and ecosystem respiration: review and improved algorithm, Glob. Change Biol., 11, 1424-1439, doi:10.1111/j.1365-2486.2005.001002.x, 2005.
92. Rodrigues, A., Pita, G., Mateus, J., Kurz-Besson, C., Casquilho, M., Cerasoli, S., Gomes, A., and Pereira, J.: Eight years of continuous carbon fluxes measurements in a Portuguese eucalypt stand under two main events: drought and felling, Agr. Forest Meteorol., 151, 493-507, doi:10.1016/j.agrformet.2010.12.007, 2011.
93. Sala, A. and Tenhunen, J.: Simulations of canopy net photosynthesis and transpiration in Quercus ilex L. under the influence of seasonal drought, Agr. Forest Meteorol., 78, 203-222, doi:10.1016/0168-1923(95)02250-3, 1996.
94. Santos, J., Corte-real, J., and Leite, S.: Atmospheric large-scale dynamics during the 2004/2005 winter drought in Portugal, Int. J. Climatol., 27, 571-586, doi:10.1002/joc.1425, 2007.
95. Santos, J. A., Woollings, T., and Pinto, J. G.: Are the winters 2010 and 2012 archetypes exhibiting extreme opposite behavior of the North Atlantic jet stream?, Mon.Weather Rev., 141, 3626-3640, doi:10.1175/MWR-D-13-00024.1, 2013.
96. Schotanus, P., Nieuwstadt, F., and Bruin, H.: Temperature measurement with a sonic anemometer and its application to heat and moisture fluxes, Bound.-Lay. Meteorol., 26, 81-93, doi:10.1007/BF00164332, 1983.
97. 2exchange of leaves of Quercus suber, Planta, 162, 193-203, doi:10.1007/BF00397440, 1984.
98. Tenhunen, J., Lange, O., Harley, P., Beyschlag, W., and Meyer, A.: Limitations due to water stress on leaf net photosynthesis of Quercus coccifera in the Portuguese evergreen scrub, Oecologia, 67, 23-30, doi:10.1007/BF00378446, 1985.
99. Tenhunen, J. D., Pearcy, R.W., and Lange, O. L.: Diurnal variations in leaf conductance and gas exchange in natural environments, in: Stomatal Function, Stanford University Press, 1987.
100. Tenhunen, J., Serra, A., Harley, P., Dougherty, R., and Reynolds, J.: Factors influencing carbon fixation and water use by mediterranean sclerophyll shrubs during summer drought, Oecologia, 82, 381-393, doi:10.1007/BF00317487, 1990.
101. Thomas, C. and Foken, T.: Re-evaluation of integral turbulence characteristics and their parameterisations, 15th Conference on Boundary Layer and Turbulence, 2002.
102. Trigo, R. M., Añel, J., Barriopedro, D., García-Herrera, R., Gimeno, L., Nieto, R., Castillo, R., Allen, M. R., and Massey, N.: The record winter drought of 2011-12 in the Iberian Peninsula, in: explaining extreme events of 2012 from a climate perspective, B. Am. Meteorol. Soc., 95, S41-S45, 2013.
103. Twine, T., Kustas, W., Norman, J., Cook, D., Houser, P., Meyers, T., Prueger, J., Starks, P., and Wesely, M.: Correcting eddy-covariance flux underestimates over a grassland, Agr. Forest Meteorol., 103, 279-300, doi:10.1016/S0168-1923(00)00123-4, 2000.
104. Unger, S., Máguas, C., Pereira, J. S., Aires, L. M., David, T. S., and Werner, C.: Partitioning carbon fluxes in a Mediterranean oak forest to disentangle changes in ecosystem sink strength during drought, Agr. Forest Meteorol., 149, 949-961, doi:10.1016/j.agrformet.2008.11.013, 2009.
105. Valentini, R., Matteucci, G., Dolman, A. J., Schulze, E.-D., Rebmann, C., Moors, E. J., Granier, A., Gross, P., Jensen, N. O., Pilegaard, K., Lindroth, A., Grelle, A., Bernhofer, C., Grünwald, T., Aubinet, M., Ceulemans, R., Kowalski, A. S., Vesala, T., Rannik, U., Berbigier, P., Loustau, D., Gudmundsson, J., Thorgeirsson, H., Ibrom, A., Morgenstern, K., Clement, R., Moncrieff, J., Montagnani, L., Minerbi, S., and Jarvis, P. G.: Respiration as the main determinant of carbon balance in European forests, Nature, 404, 861-865, doi:10.1038/35009084, 2000.
106. Vargas, R., Sonnentag, O., Abramowitz, G., Carrara, A., Chen, J., Ciais, P., Correia, A., Keenan, T., Kobayashi, H., Ourcival, J.-M., Papale, D., Pearson, D., Pereira, J., Piao, S., Rambal, S., and Baldocchi, D.: Drought influences the accuracy of simulated ecosystem fluxes: a model-data meta-analysis for Mediterranean oak woodlands, Ecosystems, 16, 749-764, doi:10.1007/s10021-013-9648-1, 2013.
107. Vaz, M., Pereira, J., Gazarini, L., David, T., David, J., Rodrigues, A., Maroco, J., and Chaves, M.: Drought-induced photosynthetic inhibition and autumn recovery in two Mediterranean oak species (Quercus ilex and Quercus suber), Tree Physiol., 30, 946-956, doi:10.1093/treephys/tpq044, 2010.
108. 2fluxes in Amazon forests: Fusion of eddy covariance data and the ORCHIDEE model, J. Geophys. Res.-Biogeo., 116, G02018, doi:10.1029/2010JG001544, 2011.
109. von Caemmerer, S.: Biochemical Models of Leaf Photosynthesis, CSIRO Publishing, 2000.
110. Wang, Y.-P., and Leuning, R.: A two-leaf model for canopy conductance, photosynthesis and partitioning of available energy I: Model description and comparison with a multi-layered model, Agr. Forest Meteorol., 91, 89-111, doi:10.1016/S0168-1923(98)00061-6, 1998.
111. Webb, E. K., Pearman, G. I., and Leuning, R.: Correction of flux measurements for density effects due to heat and water vapour transfer, Q. J. Roy. Meteor. Soc., 106, 85-100, doi:10.1002/qj.49710644707, 1980.
112. Werner, C. and Correia, O.: Photoinhibition in cork-oak leaves under stress: influence of the bark-stripping on the chlorophyll fluorescence emission in Quercus suber L., Trees, 10, 288-292, doi:10.1007/BF02340774, 1996.
113. Werner, C. and Máguas, C.: Carbon isotope discrimination as a tracer of functional traits in a mediterranean macchia plant community, Funct. Plant Biol., 37, 467-477, doi:10.1071/FP09081, 2010.
114. Werner, C., Correia, O., and Beyschlag, W.: Two different strategies of Mediterranean macchia plants to avoid photoinhibitory damage by excessive radiation levels during summer drought, Acta Oecol., 20, 15-23, doi:10.1016/S1146-609X(99)80011-3, 1999.
115. Werner, C., Ryel, R. J., Correia, O., and Beyschlag, W.: Effects of photoinhibition on whole-plant carbon gain assessed with a photosynthesis model, Plant Cell Environ., 24, 27-40, doi:10.1046/j.1365-3040.2001.00651.x, 2001.
116. Werner, C., Correia, O., and Beyschlag, W.: Characteristic patterns of chronic and dynamic photoinhibition of different functional groups in a Mediterranean ecosystem, Funct. Plant Biol., 29, 999-1011, doi:10.1071/PP01143, 2002.
117. Wilczak, J., Oncley, S., and Stage, S.: Sonic anemometer tilt correction algorithms, Bound.-Lay. Meteorol., 99, 127-150, doi:10.1023/A:1018966204465, 2001.
118. Zeppel, M. J. B., Wilks, J. V., and Lewis, J. D.: Impacts of extreme precipitation and seasonal changes in precipitation on plants, Biogeosciences, 11, 3083-3093, doi:10.5194/bg-11-3083-2014, 2014.
119. Zhou, S., Duursma, R. A., Medlyn, B. E., Kelly, J. W., and Prentice, I. C.: How should we model plant responses to drought? An analysis of stomatal and non-stomatal responses to water stress, Agr. Forest Meteorol., 182-183, 204-214, doi:10.1016/j.agrformet.2013.05.009, 2013.