Effects of a Severe Drought on Growth and Wood Anatomical Properties of Quercus Faginea
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IAWA Journal, Vol. 25 (2), 2004: 185–204 EFFECTS OF A SEVERE DROUGHT ON GROWTH AND WOOD ANATOMICAL PROPERTIES OF QUERCUS FAGINEA Leyre Corcuera1, Jesús Julio Camarero1,2 & Eustaquio Gil-Pelegrín1,* SUMMARY We studied the growth response to drought of a Quercus faginea Lam. stand in a xeric site in NE Spain, that experienced an intense defolia- tion in 1993–94. This event coincided with very low precipitation from November to February, the period when total monthly precipitation ex- ceeds evapotranspiration. We evaluated the effects of November–Febru- ary precipitation (recharge precipitation, RP) on internode length, radial growth, and wood anatomy. Quercus faginea showed reduced longitudi- nal and radial growth during the years with low RP, and most sampled trees did not produce latewood in 1993–94 but showed wide earlywood vessels. We observed the reverse for years with a high RP. Radial growth was enhanced by increased precipitation during January and May of the growth year. If severe droughts become more frequent, due to a greater climatic variability, extensive dieback of marginal Q. faginea popula- tions may be expected. Key words: Dendroecology, Mediterranean climate, oak, predicted hy- draulic conductance, xylem. INTRODUCTION Reduced radial growth has been frequently described in temperate and Mediterranean oak forests in Europe and North America (Delatour 1983; Tainter et al. 1983). The main causes pointed out to explain this are: pathogens, excessive competition among neigh- bouring trees, aging and increased susceptibility, air pollution, and climatic stress (Manion 1981). The climate can act as a predisposing factor to pathogens or as a direct cause of growth reduction (Hepting 1963; Tuset et al. 1996). In fact, droughts have been involved in the onset of reduction in oak growth (Tryon & True 1958; Becker & Lévy 1982; Tainter et al. 1990; Jenkins & Pallardy 1995), especially in xeric sites (Lévy et al. 1992). Water stress has also proved to be a predisposing factor to pathogen attacks on Mediterranean oaks (Vannini & Scarascia 1991). In Mediterranean areas, summer drought and winter cold are the main factors limiting plant growth (Mitrakos 1980). Both factors also affect the xylem structure (Fritts 1976; 1) Unidad de Recursos Forestales, Centro de Investigación y Tecnologia Agroalimentaria, Go- bierno de Aragón, Apdo. 727, 50080 Zaragoza, Spain. 2) Departament dʼEcologia, Facultat de Biologia, Universitat de Barcelona, Avda. Diagonal 645, 08028 Barcelona, Spain. *) Corresponding author: Dr. E. Gil-Pelegrin1 [E-mail: [email protected]]. Downloaded from Brill.com09/30/2021 01:25:47AM via free access 186 IAWA Journal, Vol. 25 (2), 2004 Corcuera, Camarero & Gil-Pelegrin — Drought effects in Quercus faginea 187 Schweingruber 1993, 1996). For instance, a decrease in the mean diameter of the conduits (tracheids, vessels) has been associated with a reduction of water availability in the soil (Carlquist 1975, 1977; Baas et al. 1983; Baas & Schweingruber 1987; Von Wilpert 1991; Zhang et al. 1992). However, winter precipitation is the likely underlying climatic factor in Mediterranean areas, because of its effect on soil water availability for subsequent growth in the spring. The vulnerability of xylem to cavitation and embolism is one of the main factors determining the drought tolerance of plants (Tyree & Sperry 1989; Cochard & Tyree 1990; Tyree & Ewers 1991; Tognetti et al. 1998). The theoretical hydraulic conductivity through xylem depends on the number of conduits and on their diameter. According to the Hagen-Poisseuille law, the hydraulic conductivity of a cylindrical conduit is proportional to its diameter raised to the fourth power (Tyree et al. 1994). Consequently, the wider vessels are more effective hydraulic conduits, but entail a greater risk of drought/frost-induced embolism (Sperry & Sullivan 1992; LoGullo et al. 1995; Sperry et al. 1994; Tyree et al. 1994). According to the Intergovernmental Panel on Climate Change (IPCC 2001), a decrease in precipitation, a rise of mean temperature (2–4 °C), and an increase both in the frequency and intensity of acute droughts are expected for the Mediterranean basin (Houghton & Yihui 2001). Indeed, a 20% reduction in the total precipitation and an increased frequency of anticyclones have been observed in the Central-Western Mediterranean Basin between 1951 and 1995 (Piervitali et al. 1997). In the Iberian Peninsula, the 1980–95 period was characterized by intense droughts, which affected several woody species (Font Tullot 1988). In the stand under study, a yellowing of leaves followed by an intense defoliation was observed in 1993–94, affecting both Quercus ilex subsp. ballota (Desf.) Samp. and Quercus faginea Lam. (Aït-Bachir 1998). While Q. ilex subsp. ballota appears in continental areas in N Africa and the Iberian Peninsula, Q. faginea is dominant in sub-Mediterranean forests in NE Spain between 600 and 1200 m a.s.l. (Blanco et al. 1997). Quercus faginea is a deciduous oak which forms ring-porous wood. It is usually found in sites with basic soils and summer precipitation greater than 100 mm (Ceballos & Ruiz de la Torre 1979; Jiménez et al. 1998). However, the most affected Q. faginea stands showed again a good vigour in 1996. This episode of intense defoliation in 1993–94 was also observed in other areas in NE Spain (Lloret & Siscart 1995). Hence, our main objective was to determine how the radial growth of Q. faginea responded to the intense 1993–94 drought. Specifically, we evaluated the relationship between low recharge precipitation (hereafter, RP) and growth reduction. We computed RP as the accumulated monthly precipitation from November prior to the growth year to February of the growth year, i.e. winter precipitation. This period was selected because: 1) during the RP months the total monthly precipitation is higher than the monthly evapotranspiration, which determines soil water availability for subsequent spring growth (Faci González & Martínez Cob 1991), and 2) radial increment is great in the Mediterreanean oaks during April–May (e.g., Zahner 1968). We focused on the growth and the xylem structure through a detailed study of the anatomy (number and diameter of vessels, vessel density) since these features were shown to be sensitive to the yearly variations in precipitation in other ring-porous Downloaded from Brill.com09/30/2021 01:25:47AM via free access 186 IAWA Journal, Vol. 25 (2), 2004 Corcuera, Camarero & Gil-Pelegrin — Drought effects in Quercus faginea 187 oaks such as Q. macrocarpa (Woodcock 1989a). Pure or mixed oak coppice stands (especially Q. ilex subsp. ballota, Q. pyrenaica Willd., and Q. faginea), characterized by a long history of intensive human use for fuel-wood and charcoal production, are very abundant in the Iberian Peninsula (Serrada et al. 1992). Due to the abandonment of their traditional management, the customary cutting frequency decreased in the 1940s (Cañellas et al. 1996). Nowadays, most of the stems are over-aged (30–50 yrs.), which might make them more sensitive to climatic disturbances such as severe droughts because of their lower hydraulic efficiency (Amorini et al. 1996). In addition, the over- aged coppice stands of Mediterranean oaks form a new landscape that is still poorly studied from an ecological point of view (Enjalbal et al. 1996). MATERIALS AND METHODS Study site A coppice stand dominated by Quercus faginea and Q. ilex subsp. ballota was selected in the Sierra de Santa Cruz-Cubel, Zaragoza, NE Spain (1° 39' W, 41° 07' N, 1177 m a.s.l.). The precipitation and temperature data were obtained from the Cubel-Casas Altas station located at c. 2 km from the stand (41° 06' N, 1° 38' W, 1108 m a.s.l.; period 1969–97). We also used precipitation data from the nearby Daroca station (41° 07' N, 1° 25' W, 779 m) to describe the temporal evolution of the rainfall in the area during the 20th century (1910–99 data). In the study area, the dry period in summer lasts c. 2 months, from the end of June to early September (Fig. 1). Since 1960, the following periods showed very low RP records: 1973–75, 1981–84, 1989, and 1992–95 (Fig. 1 & 2). In fact, the two lowest values for the total precipitation in January in the last 50 years in the study area were recorded in 1983 and 1993, respectively. The climate of the study area shows a transition from sub-Mediterranean to Medi- terranean (Allué Andrade 1990). The landscape was previously dominated by coppice stands of Q. faginea, but Q. ilex subsp. ballota is currently the most abundant tree species due to selective logging. Intense coppice management for fuelwood was carried out 40–50 years ago. The study site is located on very poor soils developed over Tertiary limestone outcrops. We assume that the thin soil and the high elevation of the study site make the trees of both species very susceptible to climatic stress (high sensitivity). Sampling procedures The sampling was done in January 1998. Ten stems were cut at mid-height (c. 1.3 m) from the S-SW side of ten dominant multistemmed Q. faginea individuals (one stem per individual) for the analyses of wood anatomical variables (n = 10). Five additional stems were sampled from other individuals for the measurement of tree-ring width, including earlywood and latewood width, and internode length (n = 15). The stems had a similar diameter and age (mean age (± SE) = 27 ± 2 years). Although the sample size was close to the minimum required in standard dendroecological studies (Fritts 1976), the detailed analysis of wood anatomical features made this the largest sample size that could be studied. The middle of the older internodal segment of each stem was transversally sectioned with a sliding microtome (Anglia Scientific AS200, UK). Downloaded from Brill.com09/30/2021 01:25:47AM via free access 188 IAWA Journal, Vol. 25 (2), 2004 Corcuera, Camarero & Gil-Pelegrin — Drought effects in Quercus faginea 189 Cubel-Casas Altas (1108 m) A 1969–1997 [29] 11.3 °C 481 mm 100 40 80 60 T (°C) T 20 40 (mm) P 20 0 0 J F M A M J J A S O N D B Daroca Cubel-Casas Altas 250 800 200 600 150 400 100 200 50 Annual precipitation (mm) Recharge precipitation (mm) Recharge 0 0 1970 1975 1980 1985 1990 1995 Time (years) Fig.