Preliminary research report

Dendrochronologia 19 (2) - 2001:

Growth of Pinus pinea and Pinus halepensis as affected by dryness, marine spray and land use changes in a Mediterranean semiarid ecosystem
Jos Ravents*, M. De Lus*, M.J Gras*, Katarina Cufar**, J.C. Gonzlez-Hidalgo***, A. Bonet*, J.R. Snchez*
* Departamento de Ecologa, Universidad de Alicante - Spain **Department of Wood Science and Technology, Biotechnical Faculty, University of Ljubljana - Slovenia *** Departamento de Geograa y Ordenacin del Territorio, Universidad de Zaragoza - Spain

Abstract Pine afforestations located on sand dunes, are among the most threatened coastal woodlands of semiarid western Mediterranean areas. In the recent time, dryness and land use changes seem to have caused a considerable degradation of these ecosystems. The marine spray affects their development as well. Our objective was to examine the possible effects of these ecological factors on tree-ring variation of Pinus pinea L. and Pinus halepensis Mill. We selected altogether 30 trees of both species from a sand dune stand located in the Guardamar Pine Woodland, Alicante, Spain. For both species we selected ve trees per each of three canopy damage levels and analysed four cores per tree. The results of dendrochronological analysis show that there exists a correlation between tree-ring widths and September-June precipitation and it is an inverse relationship between the growth-climate correlation and the degree of canopy damage. Moreover, trees showed an increased number of missing-rings during the 1990s which was strongly related to severity of canopy damage due to marine spray. Synergistic effect of defoliation produced by the marine spray and dryer conditions due to limited water availability seem to have strong affect on variation of tree-ring widths. Keywords: dendroecology; tree-ring records, dune ecosystem, missing rings, canopy damages, rainfall series

Introduction In the Western Mediterranean areas, the rainfall variability in space and time is one of the most relevant characteristics (Romero et al. 1998). For these areas, where water availability is the main limiting factor for forest development, General Circulation Models (GCMs) predict a signicant decrease in rainfall for next decades. Two recent studies based on historical rainfall records seem to agree with this prediction. Annual rainfall during 1961-1990 has signicantly decreased and interannual variability has signicantly increased (De Lus et al. 2000). Moreover, a signicant change in seasonal distribution has been detected (Gonzlez-Hidalgo et al. 2001). All this together allowed Lavorel et al. (1998) to hypothesise that Mediterranean regions, as transitional climate zones, are areas where climatic changes may have the greatest effects.
Corresponding author: J. Ravents - e-mail: jraventos@ua.es

In this context, pine afforestations located in coastal areas, belong to the most threatened woodlands of semiarid western Mediterranean areas (Barnes et al. 2000). The main species used in afforestations in Mediterranean semiarid dune ecosystems are Pinus pinea L. and Pinus halepensis Miller. These pine species are not native in the dune systems (Bols 1967), and have to survive in hard environmental conditions, with a limited water resource and exposed to a constant threatening effect of marine spray (Garrec 1994). The Guardamar Pine Woodland is a good example of this kind of afforested dune ecosystem. This area, located in the Valencian Region (SE Spain), is characterised by a semiarid climate and a clear gradient of canopy damage due to marine spray from coastal to inland areas (Gras et al. 2000). Moreover, the conservation of this ecosystem is highly

 Ravents, De Lus, Gras, Cufar, Gonzlez-Hidalgo, Bonet and Snchez

Fig. 1 - Location of the sampling site Guardamar Pine Woodland, Valencian Region, SE Spain.

related to the dynamics of the nearby Segura River (Aldeguer et al. 1997). Furthermore, dryness (De Lus 2000) and changes in water and land uses could have affected water supply to woodland. The objective of this study was to produce a preliminary chronology for these pine species in relation to its degree of canopy damage. Thereafter, we related them with precipitation series and discuss our results in relation to ideas exposed above. Materials and Methods Study area The Guardamar Pine Woodland is located in the Valencian Region (SE Spain) (Fig. 1). This area is
30 60

characterised by a semiarid climate with a mean annual rainfall of 304 mm, concentrated mainly in autumnal season. Mean annual temperatures range from 12C in the coldest month January, to 23C in the warmest August (Fig. 2). Between 1900 and 1920 the sand dunes were afforested with two pine species (Pinus pinea and Pinus halepensis) to x the dune along the coastal line (Mira 1906). Presently, forest density is about 500 ind/ha, composed of approximately 45% of P. pinea and 55% of P. halepensis with wide range of sizes (DBH from 15 cm to more than 55 cm) (Gras et al. 2000). Experimental Design During the rst months of 2000, samples of Pinus pinea (PIPI) and Pinus halepensis (PIHA) were collected from different microtopographies and sea expositions (Tab. 1). Trees were classied according to degree of marine spray injury using a three-level scale: healthy, moderately damaged (25-60% of the canopy injured), and severely defoliated (60-99% of the canopy injured) (CEC-CEPE, 1996) (Fig. 3). From each species and level of injury, ve trees were selected and four cores were extracted from each of them on the north, south, east, and west direction. The cores were sampled at breast height with an increment borer. Then, the cores were mounted, dried, and ne-sanded. After that,

25 0 20

Temperature Rainfall

50

40

15

30

10

20

10

0 J F M A My Jn Jl Ag S O N D

Month

Fig. 2 - Climate diagrams for Guardamar. Temperature in (C) (thin line) and rainfall (mm) (bold line).

Dendrochronologia 19 (2) - 2001:

Fig. 3 - Different degree of canopy damage in this Pinus spp. (a) Healthy Pinus pinea. (b) Moderately damaged Pinus halepensis, (c) Severely damaged Pinus halepensis. Notice the typical ag shape of the crown.

 Ravents, De Lus, Gras, Cufar, Gonzlez-Hidalgo, Bonet and Snchez

Tree Species

Damage

Time Span Chronology Name earliest latest PIPI_1 PIPI_2 PIPI_3 PIHA_1 PIHA_2 PIHA_3 1918 1918 1921 1927 1911 1913 1999 1999 1999 1999 1999 1998

Lenght (years) 82 82 79 73 89 87

Number of Trees 5 5 5 5 5 5

Total No. of Cores Sampled 20 20 20 20 20 20

Total No. of Cores Used in Chronologies 17 18 10 13 14 7

Pinus pinea

Healthy Moderate Severe

Pinus halepensis Healthy Moderate Severe

Tab. 1 - Basic data on tree-ring chronologies from Guardamar. The site is located at Latitude 0 41 09 W, Longitude 38 4 47 N, Elevation 5 m.s.s.l..

the ring-widths were measured to the nearest 0.01 mm using a LINTAB (Rinn 1996) measuring table. Tree-ring analysis The tree-ring series were visually cross-dated. The cross-dating was veried statistically by using the TSAP and the COFECHA programmes (Rinn 1996; Holmes 1994). The ring-width series containing less than 50 years were not considered in analysis. The cores where we could not condently assign every ring to a specic calendar year, were not used to construct a chronology. The ring-width series of individual trees were standardised using the ARSTAN programme (Cook, Peters 1981; Holmes 1994). The long-term trend was removed from each time series of ring width measurements by tting a curve and calculating an

index dened as actual ring-width for each year divided by the curve-t value. This allowed us to remove the non-climatic age and size trend, and the effects of stand dynamics (Cook, Kairiukstis 1990). The rst step stage de-trending used a modied negative exponential curve. At the second stage, a more exible t of the curve to the data was achieved by using a cubic smoothing spline, designed to remove variability on the time scales of 50 years or more. Finally, each individual series was combined into a single non autoregressive chronology by a technique known as biweight robust estimate of the mean designed to reduce the inuence of isolated outlayer values (Cook 1985) (Tab. 2). Rainfall Data Monthly rainfall data between 1950-1999 were

Total Chronology

Common Intervals Variance in rst principal component (%) 51 49 44 60 45 31

Agreement Mean Chronology Mean Standard Time span Total n with population correlation name sensitivity deviation Skewness Kurtosis (AD) of years chronology among radii PIPI-1 PIPI-2 PIPI-3 PIHA-1 PIHA-2 PIHA-3 0.24 0.24 0.22 0.28 0.28 0.25 0.29 0.32 0.29 0.32 0.32 0.25 0.44 0.91 0.47 0.23 -0.33 -0.45 0.02 1.06 0.39 -0.53 0.86 0.08 1939-1999 1936-1999 1943-1993 1942-1990 1932-1991 1934-1987 0.94 0.92 0.85 0.93 0.73 0.59 61 55 51 49 60 54 0.47 0.44 0.36 0.55 0.25 0.17

Tab. 2 - Summary statistics for the six chronologies from computer program ARSTAN.

Dendrochronologia 19 (2) - 2001:

collected from the Guardamar weather station (0 41 09 W, 38 4 47N, 5 m.s.s.l., code 7261-O Meteorological National Agency, INM). Monthly and seasonal data were checked for inhomogenities with Alexandersson test (Alexandersson 1986; Peterson et al. 1998). Results Comparison of Chronologies The tree-ring index variations for three groups

of canopy damages in both species are shown in Figs. 4 and 5. The average correlation coefcient among the trees from each species and canopy damages demonstrated similarities in growth characteristics. The correlation coefcients for P. pinea chronologies were 0.89, (healthy vs moderate); 0.72, (healthy vs severe) and 0.73 (moderate vs severe). For P. halepensis chronologies, correlation coefcients were 0.68 (healthy vs moderate); 0.67 (healthy vs severe) and 0.62 (moderate vs severe). All comparisons were signicant at p 0.001.

2.0 1.5 1.0 0.5 0.0 2.0 1.5 1.0 0.5 0.0 2.0

(a) Pinus pinea (HEALTHY)

(b)

Pinus pinea (MODERATE)

(c) 1.5 1.0 0.5 0.0

Pinus pinea (SEVERE)

1920

1930

1940

1950

1960

1970

1980

1990

2000

Year
Fig. 4 - Tree-ring index variations for Pinus pinea with different degree of canopy damage.

 Ravents, De Lus, Gras, Cufar, Gonzlez-Hidalgo, Bonet and Snchez

Tree Rings and Rainfall Both studied species showed a high sensitivity to climatic variations in mean sensitivity sensu Douglas (1936) denoted by S for Mediterranean species. In P. pinea, S value was 0.24 and in P. halepensis 0.28. The correlation between tree-ring indices (TRI) and normalised standard deviation of SeptemberJune precipitation (i.e. departure) are shown in Figs. 6 and 7. In ve cases a highly to moderately correlation was found between TRI and September-June

precipitation (Tab. 3). There was an inverse relationship between the growth-climate correlation and the degree of canopy damage. This was true for both species with an exception of moderately damaged P. pinea. Missing Rings and Damages Although tree-ring index variation of P. pinea and P. halepensis did not show any abrupt changes in the period of 1950-1990, occurrence of missingrings was mainly concentrated in the last years.

2.0 1.5 1.0 0.5 0.0 2.0 1.5 1.0 0.5 0.0 2.0 1.5 1.0 0.5 0.0

(a) Pinus halepensis (HEALTHY)

(b)

Pinus halepensis (MODERATE)

(c)

Pinus halepensis (SEVERE)

1920

1930

1940

1950

1960

1970

1980

1990

2000

Year
Fig. 5 - Tree-ring index variations for Pinus halepensis with different degree of canopy damage.

Dendrochronologia 19 (2) - 2001:

PIPI-1 R p 0.48** 0.000

PIPI-2 0.18 0.103

PIPI-3 0.36** 0.008

PIHA-1 0.57** 0.000

PIHA-2 0.46** 0.000

PIHA-3 0.30* 0.018

Tab. 3 - Relationship between chronologies of Pinus pinea and Pinus halepensis and precipitation. R = Coefcient of Correlation; p = probability value; N = 49.

Fig. 6 - The relationship between tree-ring growth and precipitation in Guardamar. (a) The relationship between Pinus pinea index for healthy trees and September-June precipitation; (b), (c) the same relationship for moderately and severely damaged Pinus pinea trees. All series are normalised.

Fig. 7 - The relationship between tree-ring growth and precipitation in Guardamar. (a) The relationship between Pinus halepensis index for healthy trees and SeptemberJune precipitation; (b), (c) the same relationship for moderately and severely damaged Pinus halepensis trees. All series are normalised.

 Ravents, De Lus, Gras, Cufar, Gonzlez-Hidalgo, Bonet and Snchez

Mean number of missing rings per tree and decade

Mean number of missing rings per tree on relation to the Aspect WE 0 2 2 1 0 1 EW 0 1.2 5 2.5 2.4 7.7 SN 0 0.2 2.3 3.3 2 10 NS 0 1.8 2 3 5.5 1

Cores with missing rings (%)

50-59 PIPI_1 PIPI_2 PIPI_3 PIHA_1 PIHA_2 PIHA_3 0 0 0 0 0 0

60-69 0 0 0 0 0 0

70-79 0 0 0 0 0 0

80-89 0 0 0 0 0 0.3

90-99 0 1.2 2.4 3.5 2.9 4.9

0 28 70 46 57 100

Tab. 4 - Statistics Missing rings in Pinus pinea and Pinus halepensis for different canopy damage.

The Tab. 4 shows the statistics on missing rings in the 1950-1999 period. In both species we observed no missing-rings from 1950 to 1988, but the trees showed an increased number of missingrings during the 1990s. Their frequency was greater in P. halepensis. It increased in the last years of the decade (Fig. 8, Tab. 4) and was strongly correlated with severity of canopy damage due to marine spray. While healthy trees in average had 0 and 3.5 missing rings, the number was signicantly higher in severely damaged trees (2.4 and 4.9 in P. pinea and P. halepensis, respectively). The frequency of missing rings was not related to the core aspect (F=0.83, p=0.494). Discussion and Conclusions The Mediterranean regions are complex landscapes where high recurrences of re, extreme rainfall events and human activities used to be the main ecological and evolutionary forces. This is especially true in the Valencian Region (East Spain) where strong changes in land use have occurred in the last 70 years. Previous studies (Richter, Eckstein 1990; Richter et al. 1991) in Pinus spp. in mountain forest sites showed a widespread common climatic signal in tree-ring widths of the Southern Spain. In coastal areas, the two investigated species show also a high sensitivity to rainfall variations. Boreux et al. (1988) reported similar results for P. pinea from the Provence Region in France (S = 0.26).

The high correlation between tree growth (with independence of damage class) and rainfall pointed out that water availability is the main factor inuencing tree growth of both pine species in semiarid dune Mediterranean conditions. Touchan and Hughes (1999) who intended to

Fig. 8 - Presence of missing rings in cores in the last 15 years. (a) Pinus pinea, (b) Pinus halepensis.

Dendrochronologia 19 (2) - 2001:

make a P. halepensis chronology in Jordania where weather conditions are similar, report on generally high frequency of missing rings. In Guardamar Pine Woodland, despite persistently high dryness, the trees did not show any missing rings between 1950 and 1988 but from 90s, a increasing number of missing rings has been found. Probably on the early decades, a extra water supply from the Segura catchtment was available. On the last decade, water demand for human activities (tourism, agriculture, high rate of urbanisation) has increased greatly on this area and could affected the availability of water for the pine woodland. Moreover, the Guardamar Pine Woodland is affected harmfully by chemical components of the marine spray, as reported in other parts of the Mediterranean coast of Spain (Diamantopoulos et al. 2000) and Italy (Bussotti et al. 1995). Synergistic effect of both factors (defoliation produced by marine spray and lower water availability) seems to have strongly affected woodland persistence. Presently, P. pinea, and particularly P. halepensis are the most representative tree species of the Spanish Mediterranean coastal landscapes. These species seem to have a high potential to study the relationship between effects of climate, water supply and damage on tree growth in the region. We intend to address the aforementioned needs in our future research. Acknowledgements
This work has been supported by a FEDER European project (1FD97-1117-C05-01) and by a CICYT Spanish project (CLI99-0957). Jos Ravents wants to thank the Sabbatical Program of the Universidad de Alicante that provided funds to stay at the LTRR Lab. (University of Arizona, Tucson) and at the Dept, of Wood Science and Technology (University of Ljubljana, Slovenia).

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