Blanco et al.

(2006) 1/35

Post-print of Forest Ecology and Management 237 (2006) 342–352.

DOI: 10.1016/j.foreco.2006.09.057

Title:

Influence of site characteristics and thinning intensity on litterfall production in two

Pinus sylvestris L. forests in the Western Pyrenees

Names of authors:

Juan A. Blanco1, 2, J. Bosco Imbert1, Federico J. Castillo1 *

Postal addresses:
1
Dpto. Ciencias del Medio Natural, Universidad Pública de Navarra, Campus de Arrosadía s/n,

Pamplona, 31006 Navarra, Spain.

2
Present address: Department of Forest Sciences, Forest Sciences Centre, University of British

Columbia, 3041-2424 Main Mall Vancouver, British Columbia , V6T 1Z4, Canada

*Corresponding author:

Tel: (34) 948 169 115

Fax (34) 948 168 930

E-mail: federico.castillo@unavarra.es

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Blanco et al. (2006) 2/35

Abstract

Litterfall production was studied for 30 months in two contrasting Pinus sylvestris L.

stands in the western Pyrenees managed under two low thinning intensities: Aspurz (625

m.a.s.l., Mediterranean climate) and Garde (1335 m.a.s.l., continental climate). Nine 0.12

ha plots per site were set up with three treatments three times repeated: removal of 0%

(not thinned, P0), 20% (P20) and 30% (P30) of initial basal area. Total litterfall was

significantly higher in Aspurz (5,533 kg ha-1 yr-1) than in Garde (3,986 kg ha-1 yr-1).

Maximum annual needle fall values were in September (Aspurz) or October (Aspurz and

Garde), and second order peaks were in June or July at both sites. In both sites, cones and

other pine organs (e.g., seeds, inflorescences) showed the same seasonal patterns, while

bark and branches, affected by sporadic occurrence of storms or snowfalls, followed no

clear pattern. Low thinning effects were site-dependent (Aspurz, P0>(P20=P30); Garde,

P0>P20>P30) due to between-site differences in stand structure. Thus, more dominant

and codominant trees (i.e., high litterfall producers) were fell in P30 than in P20 in Garde

relative to Aspurz. Needle fall in Aspurz showed unique significant positive correlations

with days of dryness and moisture deficit, probably reflecting its Mediterranean climate.

However, relative to Aspurz, Garde exhibited significant retarded responses of total

litterfall and needle fall to changes in air temperature, soil temperature and soil moisture

suggesting in part that Garde’s needles are more resistant to climatic stress. Furthermore,

litterfall was better predicted in Aspurz than in Garde, partly due to the greater influence

of sporadic windstorms and snowstorms in Garde. It appears that in these Pyrenean Pinus

sylvestris stands, site-dependent climatic variables have an important control on litterfall.

At the local scale, however, stand structure influences effects of thinning on litterfall.

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Blanco et al. (2006) 3/35

Key words: tree removal, aboveground biomass production, forest management, nutrient

cycling, stand structure, Scots pine

1. Introduction

To achieve sustainable forest management it is essential to understand forest processes at

spatial and temporal scales of interest. Among these processes is internal nutrient cycling,

which consists of fluxes of vital mineral elements circulating through the ecosystem

components. One of these fluxes is litterfall (leaves, buds, flowers, fruits, bark, branches,

etc.) which is generally the most important way of nutrient transfer to forest soil (Bray

and Gorham, 1964; Vitousek et al., 1995; Berg and Meentemeyer, 2001), although fine

roots turnover may have a more important role than is usually accepted (Vogt et al.,

1986). It is well established that litterfall amount that reaches forest soil is strongly

influenced by climate (Meentemeyer et al., 1982; Rawat and Singh, 1989; Arneth et al.,

1998). Hence, litterfall production is low in high latitudes where a short growing season

limits plant growth, but increases towards equatorial latitudes, where plants are able to

grow during the whole year (Albrektson, 1988). At a regional scale, differences in

litterfall production may exist between sites due to climate modification by topography

(Santamaría and Martín, 1998). This fact is especially important in the Iberian Peninsula,

where altitude gradients generate highly diverse environments within small areas. Indeed,

Iberian forests show greater needle fall biomass variability than that of European forests

of higher latitude (Pausas et al., 1994). Due to the importance of these local differences,

in this work we examine two contrasting stands in the western Pyrenees: a southern site,

corresponding to a highly productive Mediterranean forest with high decomposition rates

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and a northern site showing more continental conditions, low production and relatively

lower decomposition rates (Blanco et al., 2003).

In addition to natural influences on litterfall, silvicultural practices may represent an

important disturbance to forest development. Among other forest practices, commercial

thinning has become a common practice due to the fast increment in basal area of the

remaining trees (Valinger et al., 2000). Thinning is favored by many foresters as one of

the most rational ways to exploit a forest, but all types of forest utilization bear potential

changes in the natural development of an ecosystem (Oliver and Larson, 1996; Marshall,

2000). Thinning may cause a temporal increase in soil fertility as a consequence of

decomposition of logging residues (the assart flux phenomenon, sensu Kimmins, 2004).

However, decreases in litterfall (Huesbschmann et al. 1999) and therefore in nutrients

input to soil (Chertov et al. 1999; Blanco et al. 2006) following thinning, combined with

nutrient removal in harvested biomass, may result in reduction of soil fertility if such

practices are repeated in the long-term (Blanco et al. 2005). Hence, litterfall reduction has

been reported in pine forests with different thinning types (in Pinus palustris, Harrington

and Edwards, 1999; in P. ponderosa, Klemmedson et al., 1990; in P. sylvestris, Montero

et al., 1999, and Nadelhoffer et al., 2000). However, other studies have reported the

opposite result. For example, Carlyle (1998) did not detect any litterfall decrease after

thinning in P. radiata forests and Agren and Knetch (2001) actually described litterfall

increase in P. sylvestris and P. ponderosa forests following thinning. According to

Carlyle (1998), some of the litterfall released before thinning could have been stored

within the aboveground biomass (i.e., deposited over branches and stems or caught by

green leaves) and released to the soil after thinning. Agren and Knetch (2001) have

proposed that silvicultural works could produce mechanical damage in remnant trees

4
Blanco et al. (2006) 5/35

involving an increase of litterfall production one year after thinning, but stabilizing again

to normal values in following years. These contrasting results show that forest response to

silvicultural practices is influenced by very diverse factors and is therefore difficult to

predict. Tests in different types of forests and geoclimatic conditions are needed to better

characterize the forests’ response to the dual effect of geoclimatic and thinning factors.

For this purpose, the present study is part of a long-term research project that aims to

analyze the influence of site and management factors on forest nutrient cycling. The

objectives of this paper are (1) to characterize the production and temporal biomass

dynamics of different litterfall fractions in two P. sylvestris forests in the western

Pyrenees under contrasting geoclimatic conditions; (2) to determine the influence of

thinning on production and temporal biomass dynamics of different biomass litterfall

fractions in these contrasting stands, and (3) to assess the relationship between litterfall

biomasses and edapho-climatic variables in both stands.

2. Materials and methods

2.1. Study sites

Our experimental sites, Garde (42º48’50’’ N, 0º52’30’’W) and Aspurz (42º42’31’’N,

1º8’40’’ W), are located in the western Pyrenees, in the province of Navarre (northern

Spain). The study location in Garde represents an example of low-productive P. sylvestris

forests in Spain. It is an even-aged stand resulting from successful clearcutting by groups

carried out during the early 1960’s. Site index is 23 m at 80 years. Mean stand age was 37

years when thinning was begun. Tree density was 3230 trees ha-1, with a mean height of

11.3 m, a mean dbh of 13.8 cm and a mean basal area of 47.8 m2 ha-1. The plots are

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Blanco et al. (2006) 6/35

located at a mean altitude of 1335 m a.s.l. and have a mean slope of 40%. The forest is

developed on a Dystric Cambisol, with a mean annual precipitation of 1268 mm, a mean

temperature of 8.2 ºC and a cold wet continental climate according to Papadakis’

classification. The study location in Aspurz is one of the most productive P. sylvestris

forests in Spain, with a site index of 29 m at age 80 years. It is an even-aged stand

resulting from successful strip-like clearcutting carried out in the mid-1960’s. Mean stand

age was 32 years at the time of initiating thinning. Tree density was 4040 trees ha-1, with

a mean height of 12.8 m, a mean dbh of 11.7 cm and a mean basal area of 41.2 m2 ha-1.

The plots are located at a mean altitude of 625 m a.s.l. and have a mean slope of 7%. The

forest is developed on a Haplic Alisol, with a mean annual precipitation of 912 mm, a

mean temperature of 12.0 ºC and a cold wet Mediterranean climate according to

Papadakis’. Both stands had similar site preparation and were naturally regenerated.

Fagus sylvatica L. was relatively important in Aspurz but not in Garde, although tree

density for beech (dbh > 7.5 cm) in Aspurz was 134 trees ha-1, which only represented a

3.3 % of tree density for pine at this site, and most beeches occurred in the lower strata.

Climatic diagrams (Walter, 1963) for the last 30 years are shown in Figure 1. Aspurz’s

meteorological station is placed very close to the experimental plots (3 km, same

altitude). However, the nearest climatic station to Garde’s plots has an altitude 550 m

lower than that of the study site. To solve this issue, a correction factor of -0.55 °C every

100 m a.s.l. was applied to calculate temperatures at this site. Precipitation values from

this station were probably underestimates relative to those of Garde. However, both the

meteorological station and our research plots are located along a northwest-southeast axis

within the same basin (at a distance of approximately 5.9 km). Most storms come from

the Atlantic ocean and move along this axis and consequently seasonal precipitation

patterns appears to be similar. Climatic conditions in both locations for the study period

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Blanco et al. (2006) 7/35

are shown in Table 1. Bedrock for both sites consists of flysch of sandstones and

limestones.

2.2. Experimental design, litterfall collection and soil climate data sampling

Nine experimental rectangular plots (30 x 40 m) per location were set up by the Forest

Service of the Government of Navarre, following a complete randomized block design

according to the guidelines of the International Union of Forestry Research Organizations

(IUFRO) (Andrew, 1986). To avoid edge effects, the silvicultural treatment

corresponding to each plot was also applied within a strip of 5 to 10 m adjacent to the

plot. Thinning was carried out in August and November 1999 in Garde and Aspurz,

respectively. Three treatments with three replicates (one of each treatment per block

assigned at random) were established for each location: 1) P0: reference with no thinning;

2) P20: moderate low thinning (20% of basal area removed) with selection of crop trees,

mainly removing suppressed trees and some dominant or codominant trees with

malformed trunks; 3) P30: heavy low thinning (30% of basal area removed) with

selection of crop trees, removing suppressed and some intermediate trees, as well as some

dominant or codominant trees with malformed trunks. Logs and most branches from the

felled trees were left outside plot limits. Circular litter traps with 60 cm diameter and

three legs of 1 m length were used to collect the litterfall. A conic plastic mesh was

attached to this structure, with 1.5 mm mesh size and 0.5 m depth. Nine of these litter

traps were randomly settled per plot. Samples were collected the first week of each

month, from May 2000 to October 2002 and from June 2000 to October 2002 in Aspurz

and Garde, respectively. In Garde, litterfall in November 2001 and February 2002 was

not collected because of heavy snowfalls. Sampling plant material was air dried for 24

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Blanco et al. (2006) 8/35

hours and then was dried at 70º C until it reached a constant weight (about 72 hours).

Sampling material from each litter trap was divided into six different fractions: needles,

branches, bark, cones, other pine organs (seeds, buds, inflorescences, etc.) and

miscellaneous (litterfall from plant species other than P. sylvestris). Then, their dry

weights were determined. Standing biomass of needles and branches were calculated by

using site-specific allometric equations (described in Blanco et al., 2005). To determine

standing bark biomass, bark volume was calculated using the equations from Novo et al.

(2001), and then this volume was multiplied by a density of 0.34 ± 0.02 g cm-3

(determined by volumetry, n=12). Soil moisture in a layer 6 cm deep was measured

monthly in 4 random areas per plot with a Delta-T HH2 soil water probe sensor

(Thetaprobe ML2X). Soil temperature was measured every month in 3 random areas per

plot with a HI 98840 thermometer. Three moisture and temperature readings were taken

closely within each area, and were then averaged before statistical analyses.

2.3. Statistical analysis

Data were analyzed using repeated-measures ANOVAs with site, thinning intensity and

collection date as main factors. Measure of the efficiency of blocking (Quinn and

Keough, 2002) indicated that blocking was not needed, so blocks were not considered in

the analyses. When significant differences between levels of treatment were detected

(P<0.05), differences between thinning intensities were tested using lineal contrasts

between control and thinned plots, and then between P20 and P30. Comparisons between

years (year 1, from May 2000 to April 2001; year 2, from May 2001 to April 2002) for

litterfall data were carried out using a one-way ANOVA. Before data analyses, boxplots

and histograms were used to detect possible outliers and to determine the skewness of the

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Blanco et al. (2006) 9/35

data. Evaluation of residual plots and the Shapiro-Wilk test indicated the need to log10(x)

transform data to achieve an approximate homogeneity of variances and normal

distribution. Data of cones, branches and other pine organs had many zeros and therefore

were transformed to ranks (Quinn and Keough, 2002). Analysis results with data

transformed to ranks were compared with results from untransformed data, obtaining

similar results in both cases.

Pearson’s correlation analysis was used to investigate the relationships between the log10-

transformed amount of total litterfall and needle fall, and edapho-climatic variables with

different time lags. Thus, monthly precipitation, monthly mean air temperature, monthly

mean soil moisture, monthly mean soil temperature and monthly moisture deficit

(Thornthwaite and Mather, 1957) were determined for the same dates as litterfall, and

one, two and three months before this date. Additionally, for each of these variables but

moisture, we also used in the analyses a new parameter calculated by averaging the value

for the litterfall collection date (i.e., time lag = 0) and the previous monthly value (i.e.,

time lag = -1). Monthly number of dry days (precipitation outside canopy  1 mm) and

monthly effective temperature sum (sum of daily mean temperatures, threshold value + 5

°C) had lags of 0 and 1 months relative to the litterfall collection dates. All statistical

analyses were carried out using JMP version 5.0 (SAS, North Carolina, USA).

3. Results

3.1. Litterfall main fractions

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Blanco et al. (2006) 10/35

Needles were the most important litterfall fraction at both sites. Needles production in

Aspurz was higher than in Garde (F1,145 =22.256, P<0.001, Table 2). Needle fall

differences between years were not significant at both sites. This litterfall fraction had a

strong seasonal pattern, with maximal values in September (Aspurz) or October (Aspurz

and Garde) and a second order peak for both sites in June or July (Figure 2). Thinning

intensity had a significant effect on needle fall (F2,145 =21.777, P<0.001), although this

effect was different at each site (site x thinning intensity, F2,145 =3.755, P=0.026). In

Aspurz, P0 plots were the most productive (F1,73 =10.687, P=0.002), but there were no

differences between P20 and P30 (F1,73 =0.250, P=0.619). In Garde, P0 plots had the

highest litterfall production as well (F1,72 =23.443, P<0.001) and P20 plots were more

productive than P30 (F1,72 =16.082, P<0.001). No significant differences between sites

nor between thinning intensities within sites were detected for needles’ fallen biomass

percentage (Garde < Aspurz) or turnover (Garde > Aspurz) (Table 3). However, after

thinning, needles’ fallen biomass percentage increased in Aspurz (P20 = 18.8%; P30

=18.5%) and Garde (P20 =51.0%; P30 = 57.2%) relative to that of their respective P0

plots.

Branches were the second most important fraction at both sites (Table 2). Branch fall was

significantly higher (F1,145 =41.766, P<0.001) in Aspurz than in Garde. A lower amount

of branches were collected during the second year. This decrease was significant in

Aspurz in P0 and P30 and in Garde in P30 (Table 2). Unlike needle fall, a seasonal

pattern did not occur for branch fall (Figure 2). Significant differences between thinning

intensities were also observed (F2,145 =25.591, P<0.001). P0 plots were the most

productive at both sites (Table 2). No significant differences between sites nor between

thinning intensities within sites were detected for branches’ fallen biomass percentage or

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Blanco et al. (2006) 11/35

turnover (Garde > Aspurz) (Table 3). However, after thinning, branches’ fallen biomass

percentage increased in Aspurz (P20 = 15.6%; P30 =10.8%) and Garde (P20 =23.4%;

P30 = 19.3%) relative to that of their respective P0 plots.

The miscellaneous fraction was more important in Aspurz than in Garde (F1,148 =156.763,

P<0.001, Table 2). Like for needles, differences between years were not significant at

both sites. Seasonal evolution was strongly different at both forests (Figure 2). In Garde,

production of this fraction was quite low during the whole year, but in Aspurz, important

peaks were observed in November and December. Miscellaneous production was

significantly higher (F1,75= 9.354, P=0.003) in P0 than in the thinned plots in Aspurz, but

no significant differences between P20 and P30 (F1,75 =1.429, P=0.236) were found

(Table 2). In Garde, there were no differences between treatments (F2,75= 1.354,

P=0.278).

3.2. Litterfall minor fractions

Bark was the forth most important litterfall fraction in Aspurz, but the third in Garde.

Bark litterfall production was significantly higher in Aspurz (F1,145 =29.338, P<0.001,

Table 2). During the second year there was a significant increase of collected bark in P30

in Aspurz and at the same time there were significant decreases in Aspurz P0 and in

Garde P30 (Table 2). A clear seasonal pattern in bark fall was not found (Figure 3).

Thinning intensity had a significant effect on bark collection (F2,145 =8.765, P<0.001), but

it was qualified by a significant site x thinning intensity interaction (F2,145 =6.082,

P=0.003). P0 plots were the most productive in Aspurz (F1,73 =19.452, P<0.001), while

P20 produced less bark fall than P30 (F1,73 =7.980, P=0.006). However, there were no

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Blanco et al. (2006) 12/35

differences between P0 and thinned plots in Garde (F1,72 =2.187, P=0.144), but P30

produced less than P20 (F1,72 =6.690, P=0.012). Similar to needles and branches, no

significant differences between sites nor between thinning intensities within sites were

detected for bark’s fallen biomass percentage or turnover (Garde > Aspurz) (Table 3).

After thinning, bark’s fallen biomass percentage only increased in Garde (P20 =19.2; P30

= 29.4) relative to that of their respective P0 plots.

“Other pine organs” (seeds, buds, inflorescences) was the fifth most abundant litterfall

fraction. Again, Aspurz was more productive than Garde (F1,144 =65.744, P<0.001, Table

2). Inter-annual variation was significant in Aspurz, with decrease in the second year for

all thinning intensities. This litterfall fraction followed a clear seasonal pattern with

maximal values in spring (Figure 3). Differences between thinning intensities were

significant as well (F2,144 =7.878, P<0.001), with a significant site x thinning intensity

interaction (F2,144 =3.388, P=0.037). Production was similar in P0 and thinned plots in

Aspurz (F1,72 =1.879, P=0.175), but the production in P20 was significantly higher than in

P30 (F1,72 =12.892, P<0.001). In Garde, there was no significant effect of thinning (F1,72

=1.275, P=0.286).

Cones was the fraction that least contributed to litterfall biomass, as it comprised less

than 1% of total litterfall in both sites. However, the production of cones was

significantly higher in Aspurz (F1,145 =37.448, P<0.001, Table 2) than in Garde. During

the second year, production of cones was significantly lower at both forests. This litterfall

fraction followed a clear seasonal pattern with maximal values in spring (Figure 3). A

significant influence of thinning was not detected (F1,145 =0.299, P=0.742) (Table 2). Each

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Blanco et al. (2006) 13/35

year cone peaks of different magnitude were observed in Aspurz in June or July, but in

Garde a clear peak only occurred in July 2000 (Figure 3).

3.3. Total litterfall

The total amount of litterfall was significantly higher in Aspurz (F1,144 =154.063,

P<0.001, Table 2). A significant decrease of production was detected at both sites during

the second year except in Aspurz P20, where an increase was observed. A cyclical pattern

was observed during the year with maximal production at the beginning of autumn in

both sites and minimal production in winter in Aspurz or in spring in Garde (Figure 4).

The effect of thinning intensity was significant (F2,144 =15.660, P<0.001), but different at

each stand (site x thinning intensity, F2,144 =6.455, P=0.002) (Table 2). In Aspurz, P0

produced more litterfall than thinned plots (F1,72 =46.891, P<0.001), but there were no

differences between P20 and P30 (F1,72 =1.533, P=0.220). In Garde there were no

significant differences between P0 and the thinned plots (F1,72 =2.589, P=0.112) but there

were differences between P20 and P30 (F1,72 =4.125, P=0.046), P30 being the least

productive plots.

3.4. Relationships between litterfall production and edapho-climatic variables

Table 4 shows the Pearson’s correlation coefficients between total litterfall and needle

fall, and the edafo-climatic variables. All current and backward monthly (i.e., 0, -1, -2, -

3) values of air temperature (Aspurz), number of dry days (Aspurz and Garde) and soil

temperature (Aspurz and Garde) showed consistent positive correlations with litterfall

and needle fall production, unlike soil moisture in Aspurz which showed consistent

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Blanco et al. (2006) 14/35

negative correlations. Precipitation had no significant influence on the dependent

variables at either site. It is also worthwhile reporting that significant ( p<0.05) or

marginally significant (p=0.06) effects of number of dry days –1 and moisture deficit (0

and –1) on total litterfall and needle fall occurred in Aspurz but not in Garde. Correlation

values for all variables but precipitation were generally higher in Aspurz than in Garde,

and higher for needle fall than for total litterfall. The response of both dependent

variables to changes in air and soil temperature, and soil moisture was usually retarded in

Garde relative to Aspurz. Generally, variables that most explained litterfall were site-

dependent, and litterfall was better predicted in Aspurz than in Garde.

4. Discussion

4. 1. Between-sites and temporal differences in litterfall

Litterfall for all fractions, and therefore total litterfall, were significantly higher in Aspurz

than in Garde. In accordance with this result, site quality was also higher in Aspurz than

in Garde (Puertas, 2001), and mean turnover for needles, branches and bark was lower in

Aspurz than in Garde. In Garde, needles accounted for about 70% of the total litterfall, a

value proposed by several authors as a mean needle fall estimate on a global basis

(Meetenmeyer et al., 1982; Albrektson, 1988). However, the lower needle percentages in

Aspurz (between 50% and 60%) are closer to values reported by Gallardo et al. (1995) or

Enright (1999). This decrease was due in part to the high relative contribution of beech

leaves to total litterfall in Aspurz. Difference in latitude between Aspurz and Garde is too

small to explain between-site difference in needle fall, but difference in altitude could

have the same effect through changes in rainfall and temperature. Hence, Aspurz is

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Blanco et al. (2006) 15/35

located at the maximal litterfall production altitude range in temperate zones (Bray and

Gorham, 1964), while the lower temperatures and stronger winds in Garde could

negatively affect tree growth resulting in lower litterfall. Total litterfall and needle fall

reduction with altitude has also been reported by other researchers (e. g., Londsdale,

1988; Guerrero et al. 1998).

Our reference plots in Aspurz (5533 kg ha-1 yr-1) and Garde (3986 kg ha-1 yr-1) are among

the most litterfall-producer stands in Spain, Aspurz being the most productive site in the

Pyrenees. Bray and Gorham (1964) predicted 5500 kg ha-1 yr-1 in warm temperate forests

and 3500 kg ha-1yr-1 in cold temperate forests, values quite similar to our own results and

more close-fitting to that predicted by Meetenmeyer et al. (1982) in their litterfall world

map (3000 kg ha-1 yr-1). The predicted values by Londsdale’s equation (1988) are lower

than our results as well (3000 kg ha-1 yr-1). Total litterfall production significantly

decreased from year 1 to year 2. On one hand, needle fall did not show significant

changes between years either in Aspurz or in Garde. On the other hand, fall of branches

(Aspurz and Garde), bark (Aspurz and Garde) and other pine organs (Aspurz) was lower

the second year. Altogether, these results suggest that differences between years are

mainly due to falling of old branches and bark caused by windstorms. Cone production

also decreased but this event seems to be related to natural variations in three-year cycles

(Koski, 1991).

Needle fall exhibited maximum peaks in September – October and a second order

maximum in June-July. Similar patterns describing peaks of needle fall in September-

October have been described for P. sylvestris (Guerrero et al., 1998) and other Pinus

species (Klemmedson et al., 1990; González-Arias et al., 1998). However, it is delayed

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Blanco et al. (2006) 16/35

compared to other reports, which have described the main amount of needle fall from

August to October (in P. pinea, Hernández et al., 1992; in P. sylvestris, Pausas et al.,

1994 and Pausas, 1997; in P. pinaster, Hernández et al., 1992) or from July to September

(Montero et al., 1999; Roig et al., 2005, both in P. pinaster forests. The secondary peak

observed in June-July may result from nutrient retranslocation from old needles to

produce new needles, falling afterwards. With regard to branch fall, maximal peaks in

Garde appeared in winter perhaps due to snowfalls that could have caused branch breaks

(Figure 2). However, in Aspurz where snowfalls are much less frequent than in Garde

(Government of Navarre, 2006), branch fall did not have a clear pattern and peaks were

probably caused by storms. Bark production was irregular during the study period and it

seemed to be related to branch litterfall (Figures 2 and 3), probably because bark can

originate not only from stems but also from branches. In the case of the miscellaneous

fraction, the production rate in Garde was constant during the year. However, in Aspurz,

high production peaks were observed in autumn due to the input of beech leaves although

they were much less important than those of needle fall. In Garde, however, the presence

of species other than P. sylvestris was relatively low. In both sites the most important

percentage of cones and other pine organs was collected at the end of spring and

beginning of summer, following the pine flowering season. The observed amounts and

their relative importance were lower than in other studies dealing with P. sylvestris in the

Pyrenees (Puigdefábregas and Alvera, 1977; Pausas, 1997) and other pine species (Xu

and Hirata, 2002), but it is possible that our study period was coincident with a moment

of low flowering. Thus, P. sylvestris shows big variations between years in flowering

intensity, which is then reflected in variations in cone production, with peaks every three

years (Koski, 1991). As it has been shown in many other studies (e. g., Puigdefábregas

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Blanco et al. (2006) 17/35

and Albera, 1977; Pausas, 1997; Guerrero et al., 1998) annual pattern of total litterfall in

both sites and in all treatments was mainly determined by needle fall.

4. 2. Effects of thinning on litterfall

Thinning affected all litterfall fractions, with the exception of cones (Aspurz and Garde)

and other pine organs (Garde). A trend to decreasing litterfall with thinning intensity in

both sites was most consistent for total litterfall and the needle fraction, as reported in

other studies (Klemmedson et al. [1990] in P. ponderosa, Dames et al. [1998] in P.

patula, Kirschbaum [1999] in P. radiata). However, statistical analyses showed site-

dependent differences. Thus, in Aspurz there were differences between P0 and thinned

plots, but P20 and P30 produced similar amounts of major litterfall components (needles,

branches and miscellaneous), and therefore total litterfall was similar for both thinning

intensities. In Garde, however, significant differences were observed between P20 and

P30. Due to between-site differences in stand structure low-thinning effects were site-

dependent; thus, different percentages of tree types were felled in P30 from each site

(Puertas, 2001). In Aspurz, mainly dead and suppressed trees (i.e., the first type of trees

to be removed) were cut, and percentages of dominant and codominant trees (i.e., main

litterfall producers) felled were relatively low and similar for P20 and P30. On the other

hand, in Garde, mainly dead trees were cut, and percentages of dominant and codominant

trees felled were also relatively low but were much higher in P30 than in P20 (Puertas,

2001) , because removing dead trees was not enough to reach 30 % of initial basal area.

An explanation for this distinct tree distribution can be found in the differences in

geoclimatic conditions between forests. Hence, in Garde the harder climatic conditions

result in higher tree mortality, and light attenuation from dominant trees to suppressed

17
Blanco et al. (2006) 18/35

tress is reduced relative to that in Aspurz due to Garde’s higher slope. For this reason, in

Garde trees able to deal with low temperatures and winds always receive radiation to

produce biomass. In contrast, slope in Aspurz is small, and consequently the stratum of

dominant trees creates a continuous layer that greatly diminishes light reaching

suppressed trees. Therefore, these trees have small aboveground biomass and do not

produce a significant amount of litterfall.

Relative to the aboveground biomass of needles, branches and bark in P0, the percentage

of fallen needles in Aspurz and Garde, and of fallen branches and bark in Garde were

higher in their respective thinned plots (see Table 3). On one hand, after removing many

dead and suppressed trees in Garde and Aspurz, the remaining trees, mostly dominant and

codominant, may have considerably increased photosynthesis (Kozlowski et al., 1991)

and needle production relative to the P0 plots, resulting in relative increase of old falling

needles. On the other hand, the increase in fallen branches and bark in Garde, may be due

to enhancing wind effects after removing trees. Finally, significant effects of thinning on

the miscellaneous fraction in Aspurz (mainly beech leaves) were unexpected as foresters

had not fell beeches. Indeed, we have not found significant differences for beech cover in

the high canopy (>2 m) (unpublished results). However, understory (<2 m) unpublished

studies in Aspurz clearly show a reduction of beech cover with thinning intensity,

probably as a response of shadow-adapted saplings to increases in light, and this might

explain our above result.

4. 3. Relationships between litterfall production and edapho-climatic variables

18
Blanco et al. (2006) 19/35

Generally, litterfall and needle fall showed consistent positive correlations with air

temperature, number of dry days and soil temperature in both sites, and negative

correlations with soil moisture particularly in Aspurz. In this respect, needle fall may

increase due to higher levels of abscisic acid induced by water stress conditions

(Sundarapandian and Swamy, 1999). However, the response to changes in air

temperature, soil temperature and soil moisture was retarded in the continental site

(Garde) relative to that of the Mediterranean site (Aspurz). Colder winter temperatures, as

those of Garde, require thicker cuticles and other structural constituents to minimize

desiccation (Reich et al., 1995). Therefore, Garde´s needles may be more resistant to the

effects of low soil moisture than Aspurz’s needles. Additionally, lower temperatures at

the former site may slow down processes leading to needle fall. Only Aspurz was

significantly correlated to number of dry days (i. e., -1) and moisture deficit (i. e., 0 and -

1). Roig et al. (2005) have reported similar relations between moisture deficit and

litterfall production for P. pinaster in Central Spain, but in their study site the effect of

moisture deficit was not delayed, probably because summer moisture deficit was much

more intense than in Aspurz. Finally, higher correlations between environmental

variables and litterfall in Aspurz appear to be related to the higher frequency of snow

events and storms in Garde that mask somewhat seasonal phenological changes (i. e.,

needle fall).

5. Conclusions

In accordance with site-quality differences between sites (Aspurz > Garde) litterfall

production for all fractions was higher in Aspurz than in Garde. Our results suggest that

altitude and climate in part determine main differences in litterfall production between

19
Blanco et al. (2006) 20/35

our sites. However, climatic differences between sites and thinning appear to affect little

litterfall seasonal cycles. Low thinning effects were site-dependent (Aspurz,

P0>(P20=P30); Garde, P0>P20>P30) due to between-site differences in stand structure (i.

e., percentages of tree types). Because of these differences more dominant and

codominant trees were fell in P30 than in P20 in Garde relative to Aspurz. Consequently,

the same thinning operation (i.e., percentages of initial basal area removed) differently

affected litterfall production, and therefore C and nutrient cycling in both forests. Aspurz

showed unique significant or marginally significant correlations between total litterfall

and needle fall, and number of dry days (lag = -1) and moisture deficit (lag = 0 and -1),

probably reflecting its Mediterranean climate. The retarded responses of needle fall in

Garde relative to Aspurz suggests that Garde’s needles are more resistant to climatic

stress; lower temperatures at Garde may also slow down processes leading to needle fall.

Furthermore, litterfall was better predicted in Aspurz than in Garde, partly due to the

greater influence of sporadic windstorms and snowstorms in Garde. It appears that in

these Pyrenean Pinus sylvestris stands site-dependent climatic variables have an

important control on litterfall. At the local scale, however, stand structure influences

effects of thinning on litterfall. Therefore, these results suggest that in our region forest

management plans should be site-specific, and not extrapolated from one stand to another

without carefully assessing differences in stand structure and ecological processes.

Finally, given that our experimental site set up was not replicated, other sites with similar

characteristics should be studied to confirm the relevance of the observed patterns in the

western Pyrenees and other geographical locations.

Acknowledgments. We thank ‘Gobierno de Navarra, Departamento de Educación y

Cultura" for financial support and ‘Departamento del Medio Ambiente’ for the

20
Blanco et al. (2006) 21/35

experimental setting of silvicultural treatments and financial support. In particular, we

acknowledge Fernando Puertas, Carmen Traver and Ana Iriarte for assistance at several

stages of this work and to Carl F. Falk for his English grammar review.

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Table 1. Climatic conditions in both forest experimental sites, Aspurz and Garde, during the two years of

study (year 1, from May 2000 to April 2001; year 2, from May 2001 to April 2002).

Variable Unit Aspurz Garde

Year 1 2 1 2
-1
Rainfall mm year 615 799 1448 865
Mean air temperature ºC 12.4 11.2 8.3 9.4

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Table 2. Litterfall production (kg ha-1) at both forest sites, Aspurz and Garde separated by fractions,

treatments and localities. Year 1: May 2000 - April 2001. Year 2: May 2001 - April 2002. Mean 

standard errors are shown for the whole study period (May 2000 - October 2002). Significance in

comparisons between years, *P<0.05, ** P<0.01, ***P<0.001 (df=1, 53). Relative content (%) of each

fraction and treatment per site are also shown.

Fraction Treatment Aspurz Garde

Year 1 Year 2 Mean % Year 1 Year 2 Mean %
P0 (control) 2720 3028 3003 137.84 54 2666 2741 2774 31.83 70
Needles P20 (thinning 20%) 2428 2620 2668 85.79 58 2503 2404 2488 29.49 69
P30 (thinning 30%) 2401 2469 2568 30.50 60 2015 1873 1995 30.83 71

P0 (control) 1397 817 ** 1049 259.13 19 654 591 552 27.97 14

Branches P20 (thinning 20%) 733 592 645 62.75 14 581 614 514 12.97 14
P30 (thinning 30%) 687 523 * 575 73.40 13 666 359 * 453 137.19 16

P0 (control) 22 8 *** 17 6.56 1 19 2 *** 9 7.82 1

Cones P20 (thinning 20%) 25 7 *** 15 7.77 1 24 1 *** 11 10.18 1
P30 (thinning 30%) 24 7 *** 15 7.68 1 14 2 *** 7 5.52 1

P0 (control) 665 642 ** 611 10.14 11 404 383 364 9.52 9

Bark P20 (thinning 20%) 457 433 414 10.60 9 397 382 361 6.75 10
P30 (thinning 30%) 506 522* 483 7.46 11 304 291 * 272 5.64 10

P0 (control) 150 118 *** 142 13.96 3 46 48 68 1.08 2

Other pine
P20 (thinning 20%) 139 126 * 145 5.92 3 53 47 68 2.78 2
organs
P30 (thinning 30%) 107 81 *** 101 11.50 2 40 40 58 0.08 2

P0 (control) 687 852 645 73.52 12 227 180 188 20.98 5

Miscellaneous P20 (thinning 20%) 411 512 404 45.02 9 171 147 146 10.89 4
P30 (thinning 30%) 533 605 492 31.94 11 164 154 145 4.62 5

P0 (control) 5641 5465 *** 5533 78.71 100 4016 3945 * 3986 31.76 100
Total P20 (thinning 20%) 4193 4290 ** 4635 43.39 100 3729 3595 ** 3584 61.63 100
P30 (thinning 30%) 4258 4207 *** 4313 22.81 100 3203 2719 *** 2795 216.45 100

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Table 3. Mean ± standard error (n=3) of standing biomass (Mg ha-1), litterfall mass (Mg ha-1), percentage of standing biomass that falls per year (%) and

turnover (yr) of the three most important P. sylvestris litterfall fractions. Different letters indicate significant differences (P<0.05) with Tukey’s H.S.D.

Comparisons have been carried out for all pair-wise combinations of thinning intensities within and between sites for each row.

Fraction Variable Aspurz Garde

Control Thinning 20% Thinning 30% Control Thinning 20% Thinning 30%

Needles Standing biomass 7.79 ± 0.86 ab 5.49 ± 0.32 b 5.29 ± 0.22 b 10.07 ± 0.93 a 6.37 ± 0.36 b 5.49 ± 0.15 b

Fallen biomass 3.00 ± 0.15 a 2.67 ± 0.04 a 2.57 ± 0.07 a 2.59 ± 0.41 a 2.47 ± 0.18 a 2.20 ± 0.18 a

% fallen biomass 39.80 ± 5.66 ab 49.02 ± 3.38 a 48.81 ± 3.50 a 25.63 ± 3.37 b 38.71 ± 1.16 ab 40.29 ± 4.05 ab

Turnover 2.64 ± 0.43 ab 2.06 ± 0.15 b 2.07 ± 0.14 b 4.03 ± 0.47 a 2.59 ± 0.08 b 2.54 ± 0.28 b

Branches Standing biomass 30.11 ± 3.73 a 20.94 ± 1.11 b 17.93 ± 0.96 b 21.16 ± 1.98 b 15.82 ± 0.38 b 13.58 ± 0.11 b

Fallen biomass 1.05 ± 0.02 a 0.65 ± 0.12 ab 0.58 ± 0.04 b 0.56 ± 0.12 b 0.52 ± 0.13 b 0.44 ± 0.08 b

% fallen biomass 3.60 ± 0.46 a 3.04 ± 0.43 a 3.21 ± 0.09 a 2.69 ± 0.62 a 3.32 ± 0.89 a 3.21 ± 0.55 a

Turnover 28.86 ± 4.19 a 34.31 ± 5.20 a 31.24 ± 0.93 a 41.50 ± 9.73a 36.93 ± 12.85a 33.55 ± 6.91a

Bark Standing biomass 35.07 ± 5.42 a 24.14 ± 1.35 ab 20.29 ± 1.35 b 26.16 ± 2.44 ab 19.51 ± 0.45 b 16.75 ± 0.14 b

Fallen biomass 0.58 ± 0.01 a 0.40 ± 0.03 ab 0.46 ± 0.04 ab 0.38 ± 0.07 ab 0.34 ± 0.07 b 0.32 ± 0.02 b

% fallen biomass 1.73 ± 0.23 a 1.65 ± 0.14 a 2.27 ± 0.10 a 1.46 ± 0.32 a 1.74 ± 0.41 a 1.89 ± 0.14 a

Turnover 60.35 ± 9.31 a 61.33 ± 4.75 a 44.20 ± 1.98 a 75.66 ± 16.63 a 65.15 ± 17.11 a 53.44 ± 4.30 a

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Table 4. Pearson’s correlation coefficients (r) between edapho-climatic parameters and total and needle fall production and significance of linear

regressions. In bold case, the highest Pearson’s coefficient for each group of similar parameters. Numbers mean 0: current month; -1: previous

month; -2: two months before; -3: three months before; (0+(-1))/2: Mean value of current and previous month.

Parameter Total Litterfall Needle fall

Aspurz Garde Aspurz Garde
r P r P r P r P
Precipitation 0 0.271 0.147 0.129 0.520 0.254 0.175 0.092 0.647
Precipitation -1 -0.087 0.648 -0.180 0.368 -0.139 0.464 -0.203 0.310
Precipitation -2 0.009 0.963 -0.220 0.271 -0.131 0.489 -0.251 0.207
Precipitation -3 -0.180 0.350 -0.301 0.128 -0.078 0.688 -0.304 0.124
Precipitation (0+(-1))/2 0.013 0.944 0.116 0.565 -0.001 0.996 0.088 0.661
Air temperature 0 0.342 0.064 -0.113 0.575 0.568 0.001 0.019 0.927
Air temperature -1 0.499 0.005 0.272 0.171 0.656 <0.001 0.491 0.009
Air temperature -2 0.471 0.009 0.465 0.015 0.517 0.003 0.675 <0.001
Air temperature -3 0.370 0.048 0.281 0.155 0.319 0.091 0.491 0.009
Air temperature (0+(-1))/2 0.494 0.006 -0.343 0.080 0.701 <0.001 -0.238 0.231
Days of dryness 0 0.095 0.619 0.081 0.688 0.138 0.467 0.094 0.642
Days of dryness -1 0.355 0.059 0.139 0.499 0.477 0.009 0.220 0.280
Effective temperature 0 0.407 0.026 -0.073 0.718 0.625 <0.001 0.067 0.739
Effective temperature-1 0.583 0.001 0.227 0.265 0.726 <0.001 0.456 0.019
Moisture deficit 0 0.670 0.006 -0.102 0.730 0.674 0.006 0.014 0.961
Moisture deficit -1 0.614 0.015 0.367 0.197 0.502 0.057 0.363 0.203
Moisture deficit -2 0.167 0.585 0.096 0.779 0.028 0.928 0.081 0.813
Moisture deficit -3 -0.050 0.878 0.302 0.367 -0.188 0.383 0.285 0.395
Soil moisture 0 -0.381 0.073 -0.226 0.324 -0.528 0.010 -0.429 0.052
Soil moisture -1 -0.639 0.001 -0.119 0.617 -0.649 0.001 -0.304 0.183
Soil moisture -2 -0.642 0.002 -0.590 0.010 -0.648 0.002 0.700 0.001
Soil moisture -3 -0.514 0.020 -0.482 0.050 -0.406 0.076 -0.574 0.016
Soil moisture (0+(-1))/2 -0.551 0.008 -0.448 0.054 -0.638 0.001 0.628 0.004
Soil temperature 0 0.514 0.012 0.063 0.787 0.729 <0.001 0.340 0.132
Soil temperature -1 0.612 0.003 0.401 0.089 0.762 <0.001 0.591 0.008
Soil temperature -2 0.487 0.025 0.440 0.068 0.511 <0.018 0.634 0.005
Soil temperature -3 0.329 0.157 0.627 0.005 0.135 0.570 0.615 0.007
Soil temperature (0+(-1))/2 0.582 0.005 0.218 0.369 0.799 <0.001 0.490 0.033

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Figure captions

Figure 1. Climatic diagrams of the experimental sites during the last 30 years. Dashed line: mean monthly

precipitation; Solid line: mean monthly temperature; y: number of years considered; T: mean annual

temperature (°C); P: mean annual amount of precipitation (mm); TM : absolute maximum temperature

(°C); tM : mean daily minimum temperature of the hottest month (°C); tm: mean daily minimum

temperature of the coldest month (°C); Tm : absolute minimum temperature (°C). Oblique striped area

shows months with an absolute minimum temperature below 0 °C.

Figure 2. Evolution of the more abundant litterfall fractions, needles, branches and miscellaneous

collected during the period May 2000-October 2002 and June 2000-October 2002 in Aspurz and Garde,

respectively. Litterfall production is expressed as the mean  standard error, n=27.

Figure 3. Evolution of the less abundant litterfall fractions, cones, bark and other pine organs collected

during the period May 2000-October 2002 and June 2000-October 2002 in Aspurz and Garde,

respectively. Litterfall production is expressed as the mean  standard error, n=27.

Figure 4. Evolution of total litterfall collected during the period May 2000-October 2002 and June 2000-

October 2002 in Aspurz and Garde, respectively. Litterfall production is expressed as the mean  standard

error, n=27.

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Blanco et al. (2006) 32/35

Figure 1 (Blanco et al.)

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Blanco et al. (2006) 33/35

Figure 2 (Blanco et al.)

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Blanco et al. (2006) 34/35

Figure 3 (Blanco et al.)

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Blanco et al. (2006) 35/35

Figure 4 (Blanco et al.)

35