Agricultural and Forest Meteorology 204 (2015) 48–57
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Agricultural and Forest Meteorology
j o ur na l ho me pag e: www.elsevier.com/locate/agrformet
Effects of an extremely dry winter on net ecosystem carbon exchange
and tree phenology at a cork oak woodland
Filipe Costa-e-Silva
a,∗
, Alexandra C. Correia
a
, Arndt Piayda
b
, Maren Dubbert
c
,
Corinna Rebmann
b
, Matthias Cuntz
b
, Christiane Werner
c
, Jorge Soares David
a
,
João Santos Pereira
a
a
CEF, Instituto Superior de Agronomia, Universidade de Lisboa, Tapada da Ajuda, 1349-017 Lisboa, Portugal
b
Deptartment Computational Hydrosystems, UFZ Helmholtz Centre for Environmental Research, Permoserstraße 15, 04318 Leipzig, Germany
c
Agroecosystem Research, BayCeer, University of Bayreuth Universitätsstraße 30, 95447 Bayreuth, Germany
a r t i c l e i n f o
Article history:
Received 2 June 2014
Received in revised form 12 January 2015
Accepted 29 January 2015
Keywords:
CO2 fluxes
Evergreen oak
Leaf area index
Mediterranean woodland
Quercus suber L.
Tree diameter increment
a b s t r a c t
In seasonally dry climates, such as the Mediterranean, lack of rainfall in the usually wet winter may
originate severe droughts which are a main cause of inter-annual variation in carbon sequestration. Leaf
phenology variability may alter the seasonal pattern of photosynthetic uptake, which in turn is deter-
mined by leaf gas exchange limitations. The current study is based on the monitoring of an extremely dry
winter in an evergreen cork oak woodland under the Mediterranean climate of central Portugal. Results
are focused on net ecosystem CO
2
exchange (NEE), phenology and tree growth measurements during two
contrasting years: 2011, a wet year with a typical summer drought pattern and 2012, with an extremely
unusual dry winter (only 10 mm of total rainfall) that exacerbated the following summer drought effects.
Main aims of this study were to assess the effects of an extreme dry winter in (1) annual and seasonal net
ecosystem CO
2
exchange, and in (2) cork oak phenology. The dry year 2012 was marked by a 45% lower
carbon sequestration (−214 vs. −388 g C m
−2
year
−1
) and a 63% lower annual tree diameter growth but
only a 9% lower leaf area index compared to the wet year 2011. A significant reduction of 15% in yearly
carbon sequestration was associated with leaf phenological events of canopy renewal in the early spring.
In contrast to male flower production, fruit setting was severely depressed by water stress with a 54%
decrease during the dry year. Our results suggest that leaf growth and leaf area maintenance are resilient
ecophysiological processes under winter drought and are a priority carbon sink for photoassimilates in
contrast to tree diameter growth. Thus, carbon sequestration reductions under low water availabilities
in cork oak woodland should be ascribed to stomatal regulation or photosynthetic limitations and to a
lesser extent to leaf area reductions.
© 2015 Elsevier B.V. All rights reserved.
1. Introduction
Cork oak (Quercus suber L.) open woodlands cover an area
of about 2–2.5 million ha in the western Mediterranean (Aronson
et al., 2009). These are man-made ecosystems exploited with
Abbreviations: (C/N), carbon-to-nitrogen ratio; (DBH), diameter at breast height;
(DOY), day of year; (ET), ecosystem evapotranspiration; (GPP), gross primary pro-
ductivity; (LAI), leaf area index; (LAImax), maximum leaf area index; (LAI
min
),
minimum leaf area index; (LUE), light use efficiency; (NEE), net ecosystem exchange;
(PAR), photosynthetically active radiation; (PLC), percentage loss in hydraulic con-
ductivity; (
md
), midday leaf water potential; (
pd
), predawn leaf water potential;
(Reco), ecosystem respiration; (se), standard error of the mean; (SLA), specific leaf
area; (SWC), soil water content; (u*), friction velocity; (VPDmax), maximum vapor
pressure deficit.
∗
Corresponding author. Tel.: +351 21365 3515; fax: +351 21365 3338.
E-mail address: filipecs@isa.utl.pt (F. Costa-e-Silva).
low-impact agro-forestry, with high biodiversity and conservation
value (Bugalho et al., 2011). In Portugal these woodlands cover
0.74 million ha and represent 23% of all forested area. Cork oak
has a significant economic value. It provides 0.7% of Portugal gross
domestic product and supplies 54% of the worldwide cork pro-
duction (Evangelista, 2010). Cork is a natural product consisting
of continuous annual layers of suberized tissue produced by phel-
logen, a secondary meristem wrapping the inner bark. Cork wine
bottle stoppers is the main product and to obtain commercial grade
cork stripping is done traditionally every 9 years. Cork removal can
only be safely done when the phellogen cells are actively dividing,
in late-spring and early-summer, to prevent injuries to the tree
(Costa et al., 2003).
Cork oak is well adapted to the adverse semiarid Mediter-
ranean climate and its ecophysiology has been well studied in
the last decades (e.g., Otieno et al., 2007; Pereira et al., 2009; Vaz
http://dx.doi.org/10.1016/j.agrformet.2015.01.017
0168-1923/© 2015 Elsevier B.V. All rights reserved.