Forest Ecology and Management 250 (2007) 206214

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Forest structure and C stocks in natural Fagus sylvatica forest in

southern Europe: The effects of past management
Agustn Merino a,*, Carlos Real b, Juan G. Alvarez-Gonzalez c,
Manuel A. Rodrguez-Guitian d
a
Department of Soil Science and Agricultural Chemistry, Escuela Politecnica Superior, Universidad de Santiago de Compostela, E-27002 Lugo, Spain
b
Department of Cellular Biology and Ecology, Escuela Politecnica Superior, Universidad de Santiago de Compostela, E-27002 Lugo, Spain
c
Department of Agriculture and Forestry Engineering, Escuela Politecnica Superior, Universidad de Santiago de Compostela, E-27002 Lugo, Spain
d
Department of Crop Production, Escuela Politecnica Superior, Universidad de Santiago de Compostela, E-27002 Lugo, Spain
Received 14 November 2006; received in revised form 19 April 2007; accepted 5 May 2007

Abstract
Managed forests often differ substantially from undisturbed forests in terms of tree structure and diversity. By altering the forest structure,
management may affect the C stored in biomass and soil. A survey of 58 natural stands located in the south-westernmost limit of European beech
forests was carried out to assess how the C pools are affected by the changes in tree structural diversity resulting from past management. The mean
tree density, basal area and the number of large trees found in unmanaged forests were similar to those corresponding to virgin beech forests in
Central Europe, whereas large live trees were totally absent from partially cut stands. Analysis of the Evenness index and the Gini coefficient
indicated high structural diversity in the three stand types. The results of the KolmogorovSmirnov test used to compare the diameter distributions
of each group revealed significant differences between stand types in terms of distributions of total tree species and of Fagus sylvatica.
The mean C stocks in the whole ecosystem trees, litter layer and mineral soil ranged from 220 to 770 Mg ha1 (average 380 Mg ha1). Tree
biomass (above and belowground), which averaged 293 Mg C ha1, constituted the main C pool of the system (5097%). The statistical test
(KolmogorovSmirnov) revealed differences in the distribution of C pools in tree biomass between unmanaged and partially cut stands. As a
consequence of the presence of large trees, in some unmanaged stands the C stock in tree biomass was as high as 500600 Mg C ha1. In the
partially cut stands, most of the C was mainly accumulated in trees smaller than 20 cm dbh, whereas in unmanaged stands the 30% of tree C pool
was found in trees larger than 50 cm dbh. Furthermore, many unmanaged stands showed a larger C pool in the litter layer. The C content of
mineral soils ranged from 40 to 260 Mg C ha1 and it was especially high in umbrisols. In conclusion, the implementation of protective measures
in these fragile ecosystems may help to maintain the highly heterogeneous tree structure and enhance the role of both soils and trees as long-term
C sinks.
# 2007 Elsevier B.V. All rights reserved.

Keywords: Fagus sylvatica; Natural forests; Stand structure; Forest C pools; Soil organic matter

1. Introduction
Natural forests show a higher degree of biological diversity
than commercial forest plantations due to the heterogeneity of
their structure and composition. Fragmentation and manage-
ment of natural forests, mainly carried out in the last decades,
have led to lower species richness and to decreased variability
in tree diameter (Lindenmayer and Franklin, 2002).
* Corresponding author. Tel.: +34 982 285900; fax: +34 982 241835.
E-mail address: amerino@lugo.usc.es (A. Merino).
0378-1127/$ see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2007.05.016
Beech woods are one of the most important types of forest in
Europe, from both an ecological and a socioeconomic point of
view. Past management of these forests has led to diverse
situations ranging from natural, virgin beech forests covering
large areas in only a few countries (Albania, Romania or
Ukraine; see Meyer et al., 2003; Oheimb et al., 2005) to
intensively managed beech forests prevailing throughout most
of Europe (Koop and Hilgen, 1987; Emborg et al., 2000;
Oheimb et al., 2005). Many of these forests that are relatively
unaffected by human influence are included in protected areas
established for each country (National and Natural Parks,
Biological Reserves, etc.) or are selected as Sites of
Community Interest in recognition of their environmental
 A. Merino et al. / Forest Ecology and Management 250 (2007) 206214
value, by application of the European Union Habitats
Directive (Council Directive (92/43/EEC).
Living trees and soils in natural and semi-natural forests
constitute major long-term stocks of organic C in Europe.
However, a better understanding of how variations in the forest
structure affect C dynamics is necessary for predicting potential
losses and storage of C. Forest degradation can disrupt the C
cycle through direct effects on tree biomass (e.g. the ageclass
distribution of the forest shifts to younger stands containing less
C; Vieira et al., 2004) and on soil organic matter (e.g. less litter
production and higher rates of litter and organic matter
decomposition (see, e.g. Dirham, 1998). On the other hand,
changes in forest management methods, the abandonment of
mountainous regions and the implementation of conservation
measures in natural forests in Europe are leading to substantial
increases in the forest C stock (UN-ECE/FAO, 2000).
Nevertheless, since biomass accumulation depends on the
extent and intensity of the disturbance as well as different
environmental parameters, such as the temperature during the
growing season, soil moisture and soil fertility, the potential C
sequestration in these forests remains uncertain.
In contrast to the extensive information on certain ecological
aspects of beech forests in Central Europe (Peters, 1997), precise
data on these types of forests in southern Europe is scarce.
Furthermore, because of the lack of long-term records, the
dynamics of virgin or semi-natural beech forests is not well
known. One of the regions for which the lack of information is
most evident is the mountains of northern Spain, including the
southernmost boundary of Atlantic deciduous forest in Europe
(Peinado and Rivas-Martnez, 1987). In this area, European
beech is a climax species (Gandullo Guitierrez et al., 2004),
which makes up uneven-aged forests, characterized by natural
forest regeneration and high growing stocks. As a consequence of
past land clearance and fires, these forests have been affected by
severe deforestation and long-term fragmentation (many of the
forest patches are less than 3 ha in size; Garca et al., 2005),
whereas the forest management carried out in the past has been
very much variable. Thus, stands confined to steep slopes, where
access is difficult, have been maintained with little or no active
management. In others stands, on the contrary, low-intensity
interventions based on poorly regulated selective logging to
obtain firewood and charcoal have been carried out in the past.
In spite of the environmental value of these forest fragments,
comparative studies of plantsoil interactions in near-natural
forests and managed forests are scarce. Thus, although the
effects of past management on regeneration patterns (Rozas,
2003) and community structure (Onaindia et al., 2004) have
been investigated, no studies have addressed the heterogeneity
of tree structure in these systems. Relatively little is known
about the quantity and distribution of C stored in these forests
(Santa Regina and Tarazona, 1999) and the effect of past
management on the C reservoir is unknown.
The aim of the present study was to assess possible effects of
past management methods on forest structure and how this
affects the C pools of the system. This information will be
useful in designing future planning strategies to preserve the
environmental functions of these fragile forest ecosystems.
207
2. Materials and methods
2.1. Habitat characteristics
The study was conducted in 58 native woodland fragments
dominated by European beech (Fagus sylvatica L.), in a
mountainous area of 250 km2 located in northwestern Spain
(between 68400 68250 N and 428550 438300 W). The area
constitutes the present-day westernmost limit of beech forests
in Europe and contains several EU Sites of Community
Interest belonging to the Natura 2000 Network. The forests
are included in 9120 Beech forests with Ilex and Taxus, rich in
epiphytes (Ilici-Fagion) and in the European Unions Council
Directive/92/43/CEE Annex I, are considered as habitats of
community interest and are therefore protected. Although
European beech is the only tree species in the canopy of some
stands, other trees, such as oaks (Quercus robur L., Quercus
pyrenaica, Will.), birch (Betula alba L.), sweet chestnut
(Castanea sativa Mill.), holly (Ilex aquifolium L.), mountain
ash (Sorbus aucuparia L.) and hazel (Corylus avellana L.) are
present in many cases. From a biogeographical point of view,
these forests are distributed among three unitsNorthern
Galician-Asturian, Navian-Ancarensian and Lacian-Narceen-
sian subsectorswhich form part of the Cantabrian-Atlantic
Subprovince (Rivas-Martnez et al., 2002; Rodrguez-Guitian
et al., 2003).
The surface area covered by each sampled forest ranges
between 3 and 43 ha and is normally less than 5 ha. The area
covers a wide range of altitudes, between 160 and 1400 m a.s.l.
The climate is characterized by mild and humid winters, and
summers with a slight water deficit. The annual average
temperatures between sites vary between 6.0 and 13.5 8C and
the annual rainfall ranges between 1200 and >2000 mm. The
forests occur on a broad spectrum of soils ranging: (a) from
strongly acidic soils developed from siliceous rocks (mixtures
of schist and quartzite) to base-rich soils on limestone, and (b)
from shallow, stony and poorly developed soils of steep
terrains to deep soils on depressions developed on colluvial
deposits. Soil types are predominantly humic leptosols/
humic lithic dystrudepts, rendsic leptosols/humic eutrudepts,
dystric cambisols/typic dystrudepts and humic umbrisols/
humic dystrudepts (FAO-UNESCO, 1998; Soil Survey Staff,
1999).
2.2. Type of forests studied
Three different types of beech-dominated stands were
distinguished according to their disturbance and management
histories. Categorization of each forest was based on
information provided by local forest managers and inhabitants,
as well as visual examination (presence of stumps, gaps).
(i) Unmanaged mature stands (UM stands, 32 stands): forests
mainly located in areas of difficult access and where there
is no evidence of recent exploitation for wood, managed as
wood pastures or subject to other direct human dis-
turbance;
208 A. Merino et al. / Forest Ecology and Management 250 (2007) 206214
(ii) partially cut stands (PC stands, 21 stands): forests that have
been subject to a variety of undefined partial felling of
large-diameter trees for at least the past 50 years and but
not in the last 5 years;
(iii) abandoned pollard stands (AP stands, 5 stands), in which
crowns were removed: forests dominated by large trees
managed in the past (>50 years ago) for obtaining
firewood or charcoal.
2.3. Tree measurements, sampling and analysis
In each site, three circular sample plots (314 m2) were
established at random. All living trees were identified to species,
and their height and diameter at breast height (dbh) were
measured. Each individual in a sample plot was classified as a
tree (7.5 cm dbh) or a sapling (57.5 cm dbh). The basal area
per hectare (G) and the number of trees per hectare (N) were
calculated from the inventory data, using only the live tree data.
Samples of litter layer and mineral soils were collected in
July 2001. In each stand, six samples of litter layerincluding
leaves, twigs and reproductive organswere collected from
within randomly placed 0.3 m-diameter rings. Samples of litter
were pooled to provide two composite samples per stand. For
mineral soil horizons, three pits were dug in the 314 m2 circular
sample plots in each stand. Samples for determination of bulk
density were taken from each horizon, with a brass core (40 mm
long, 55 mm inner diameter). Soil samples for determination of
coarse fragments (i.e. particles >2 mm) and for C analyses
were air-dried and sieved with a 2 mm screen. Total C was
analyzed in litter and mineral soils samples with an Elemental
Analyzer (LECO).
2.4. C contents in soils and biomass
Total C storage in each soil horizon was calculated by
considering the mean values of C concentration, depth and bulk
density and the value adjusted for gravel content for each
horizon. For litter layer, C contents were calculated by
multiplying the concentrations of C by the average weight of
the layer. These procedures have been described in previous
studies (i.e. Balboa et al., 2006).
The aboveground tree biomass (>7.5 cm dbh) was estimated
for UM and PC stands using allometric equations previously
published for the main species: Fagus syvaltica and C. sativa
(Montero et al., 2006), Q. robur (Balboa et al., 2006) and B.
alba (Balboa et al., 2005). The model proposed by Le Goff and
Ottorini (2001) was applied for root biomass. Since the
allometric equations are probably not suitable for pollard trees,
they were not used for AP stands. For all vegetation
compartments, a carbon concentration of 50% (dry matter
basis) was assumed (Matthews, 2003).
2.5. Data analysis
2.5.1. Stand structure
The structural diversity of the three stand types was
characterized by using two different indices: the Weighted
Evenness index (WEv) and the Gini coefficient (Weiner and
Solbrig, 1984). The indices were calculated as follows:
X k
h pi log2 h pi =k hi
WEv  (1)
k hi 2=k log2 k
i1
 
Pn Pn  

i1 d
j1  i  d j
Gini (2)
2 n n  1 d
where pi is the proportion of trees in the diameter class i; k the
number of diameter classes; h the mean height of the stand; hi
the mean height of the diameter class i; n the number of trees of
the stand and d is the mean diameter of the stand.
The Weighted Evenness index is a modified version of the
commonly used Evenness index, in which the height values are
included, as the structural diversity of a stand depends on both
diameter and height distributions (Weber, 2000). The value of
this index increases when the number of diameter classes and
the heterogeneity of heights increase. The minimum value of
the Gini coefficient is 0 when the diameter of all trees is the
same, and the theoretical maximum value is 1, in an infinite
population in which the diameter of all individuals except one,
is 0. Both of these indices have been widely used in structural
diversity studies (e.g. Knox et al., 1989; Rouvinen and
Kuuluvainen, 2005).
The empirical diameter distribution of each stand was
described using the Weibull probability density function, one of
the most commonly used functions in sizeclass modelling
because of its flexibility and simplicity (Maltamo et al., 1995).
The mathematical expression of the three-parameter Weibull
density function is as follows:
  
c x  a c1 xa=bc
f x e (3)
b b
where x is the random variable, a is the location parameter that
defines the origin of the function, b is the scale parameter and c
is the shape parameter that controls the skewness.
The Weibull parameters were estimated using the parameter
recovery approach of the moments. In this method, the
parameters of the Weibull function are recovered from the first
two order moments of the diameter distribution (i.e. the mean
and variance, respectively), assuming that the location
parameter (a) is equal to the minimum diameter. This approach
simplifies the model and provides similar results to the three-
parameter Weibull function (Maltamo et al., 1995).
The KruskalWallis non-parametric test was used to
compare the structural diversity indices and the estimated
Weibull parameters among the three stand types.
A more detailed comparison of the diameter distributions
(all the stands of the same type together) was based on the
KolmogorovSmirnov test for goodness-of-fit. Since the results
of this test only are relevant if the data sets are balanced, the
pooled distributions (all the data of the same stand type
together) were used.
 A. Merino et al. / Forest Ecology and Management 250 (2007) 206214 209

Finally, to describe the shape of the dbh-distributions, the
shape index proposed by Siipilehto (1999) was calculated for
the pooled diameter distributions and separately for F. sylvatica
and other tree species. The basal area of the basal-area-
weighted median tree (m2) was multiplied by the stem number
(ha1) to obtain the calculated stand basal area. The shape
index is then calculated by dividing the observed stand basal
area (m2 ha1) by the calculated stand basal area. This index
has the following properties (Rouvinen and Kuuluvainen,
2005): (i) if the dbh-distribution resembles a peaked unimodal
distribution, the value of the index is approximately 1; (ii) the
index values of unimodal distributions range from 0.54 to 1
depending on the deviation and the skewness in the diameter
distribution (iii) when distribution is uniform in shape or
resembles a typical reverse-J shape, the index values decrease
to approximately 0.54 and 0.48, respectively.
2.5.2. Other statistical procedures
Kernel-smoothing techniques (Wand and Jones, 1995) were
used for estimating a hypothesized probability density function
from observed carbon content in the different C pools
considered in this study: tree biomass, litter layer and mineral
soil. Kernel density estimation is a non-parametric technique
for density estimation in which a known density function (the
kernel) is averaged across the observed data points to create a
smooth approximation. This method is more efficient than
histograms or box-and-whisker plots in creating a graphical
representation of the data distribution. Multimodality, skew-
ness, outliers and other characteristics of the distributions can
be easily seen in the resultant graphs. A Gaussian kernel was
selected and the direct plug-in method with two levels of
functional estimation was used to compute the bandwidth
(Sheather and Jones, 1991). The KernSmooth package (Ripley,
2002), running under the R environment (R Development Core
Team, 2004), was used to apply this technique to the data. To
compare the data distribution of C content between unmanaged
and partially cut stands or among soil types we employed the
KolmogorovSmirnov two-sample test.
3. Results
3.1. Stand characteristics and dbh-distributions
The number of tree species did not differ significantly
among the three types of forests (Table 1). The PC stands
tended to be comprised of more stems per hectare than the other
two stand types, but with a lower basal area (Table 1).
The curves showing the cumulative percentage of basal area
per diameter class (Fig. 1) indicate that the accumulation of
basal area is more gradual in AP stands and UM stands due to
the presence of old, large trees in these types of stand. The
greatest differences in number of stems per hectare for the three
stand types were found in smaller diameter classes (7.5 cm).
The PC stands showed the highest number of small diameter
trees, i.e. more than twice those found in the other stand types,
probably due to the growth of saplings after extraction of
dominant trees.
The values of the structural diversity indices are shown in
Table 2 for each stand type considering all species, F. sylvatica
only, or all species, except the latter. The mean value of the
Weighted Evenness index for all the species ranged from 0.74
to 0.80, indicating high structural diversity in the three types of
stand. The value of this index for diameter classes of all species
was significantly lower for PC stands than for the other two
stand types due to the absence of old, larger trees in the latter.
The Gini coefficient was always significantly higher for AP
stands due to the more gradual accumulation of number of trees
per diameter class than in the other two types of stands.
The pooled tree diameter distributions of the three stand
types are shown in Fig. 2. The UM and PC stands appeared to

Table 1
Stand characteristics of the three types of forest stands for pooled tree data (i.e. all species), Fagus sylvatica and all species except the latter
Unmanaged stands (UM) Partially cut stands (PC) Abandoned pollard stands (AP)
Species richness 2.86 (1.61) 1.90 (1.53) 3.20 (0.83)
1
Trees (>7.5 cm) ha 568.9 (238.4) 758.3 (213.4) 445.9 (98.7)
Fagus sylvatica 432.8 (187.3) 676.7 (245.3) 312.1 (88.7)
Other species 136.1 (141.5) 81.6 (88.6) 133.8 (51.0)
Basal area (m2 ha1) 48.0 (19.4) 37.6 (10.7)) 104.0 (28.1)
Fagus sylvatica 42.9 (20.5) 34.0 (10.0) 89.6 (37.8)
Other species 5.1 (4.5) 3.6 (3.6) 14.4 (17.4)
Mean diameter (cm) 29.9 (7.9) 23.0 (4.5) 43.4 (4.8)
Fagus sylvatica 33.7 24.3 (5.3) 52.0 (14.7)
Other species 19.8 (7.8) 16.1 (6.0) 26.8 (14.8)
Mean height (m) 18.7 (4.0) 15.7 (2.5) 17.6 (1.7)
Fagus sylvatica 20.1 (4.4) 16.2 (2.4) 19.5 (1.4)
Other species 14.5 (5.1) 12.1 (3.2) 13.4 (5.4)
Saplings (57.5 cm) ha1 1522.8 (403.6) 3197.8 (474.6) 1502.9 (278.7)
Fagus sylvatica 288.1 (58.7) 1345.1 (349.2) 165.6 (40.4)
Other species 1234.7 (371.0) 1852.7 (374.5) 1337.3 (261.4)
The values are means and standard deviations (in brackets).
210 A. Merino et al. / Forest Ecology and Management 250 (2007) 206214

3.2. Carbon pools in tree biomass and soils

The C contents, as well as the contribution of the living

Fig. 1. Curves of cumulative percentage of basal area for diameter classes (cm),
for the pooled data of the three stand types.
exhibit a reverse-J shape distribution, with a non-linear
reduction in number of trees per hectare with increasing
diameter class. However, the pooled distribution of AP stands
exhibited a linear decrease or uniform distribution in tree
densities across diameter classes (15 cm) due to the specific
management objective of this type of stand.
The results of the KolmogorovSmirnov test (Table 3)
indicated that the three-pooled distributions differed signifi-
cantly for both total tree species and F. sylvatica distributions.
The mean, minimum, maximum and standard deviation of the
predicted scale (b) and shape (c) Weibull parameters for each
stand type are shown in Table 4. The scale parameter differed
significantly (KruskalWallis test, p < 0.05) among stand types
for both total tree and beech distributions, while the shape
parameter was very homogeneous (only differences for PC
stands compared with AP stands). The values of the shape
parameter were always lower than 1 according to the
descending patterns of the observed diameter distributions.
Analysis of the mean values of the shape index provided similar
results (Siipilehto, 1999). This index ranged from 0.47 to 0.49
for all the species together, which reflects the reverse-J shape
of the distributions (Table 2).
and the non-living compartments, varied substantially with
the type of soil and the type of forest (Fig. 3). The total C pool in
the ecosystems ranged from 220 to 770 Mg ha1 (mean,
673 Mg ha1).
Tree biomass (above and belowground) averaged
293 Mg C ha1 and constituted the main C pool of the system
(5097%). Fig. 4 shows the kernel-smoothed distribution
functions estimated for the tree C pool in two types of stands
considered. The hypothesized probability density function of
unmanaged stands and partially cut stands are clearly different
for total tree biomass (Fig. 4a), aboveground biomass (Fig. 4b)
and litter layer (Fig. 4c). These differences are associated to
differences in stand structure and were pointed out for the
results of the KolmogorovSmirnov test at a 5% of significance
level for the three carbon pools analysed. Unmanaged stands
showed higher values of C accumulation in tree biomass due to
the importance of old, large trees. In these stands more than
30% of the basal area is accumulated in trees larger than
50 cm dbh (Fig. 1) where in partially cut stands these trees
represents only the 3% of the basal area. As a consequence,
some of the unmanaged stands showed tree C stocks as high as
500600 Mg C ha1 (Fig. 3).
The contribution of litter to the C pool of the system was less
than 5% in all cases. In PC stands the mass of litter organic
material was linearly related to basal area (r = 0.40, p < 0.05).
This parameter did not follow any clear trend with altitude for
either of the forest types. This C pool followed a different
distribution in the two types of stands (probability value for the
KolmogorovSmirnov test, p = 0.019), which reflects a higher
litter production in the unmanaged stands (Fig. 4c).
The amount of C stored in the A soil horizon of the UM
stands was slightly related to the amount of litter layer
(r = 0.47, p < 0.05) and altitude (r = 0.34, p < 0.05). In the PC
stands, a slight relationship between C content and altitude was
found (r = 0.45, p < 0.05). The C contents in the upper mineral
soil (A or A1 horizon) did not differ among types of forest or
type of soil. The amount of C stored in the whole soil profile
depended on the gravel content (r = 0.64, p < 0.01), soil depth

Table 2
Structural diversity indices and shape index of the three types of forest stands for pooled tree data, Fagus sylvatica and all species except the latter
Unmanaged (UM) Partially cut (PC) Abandoned pollard (AP)
Weighted Evenness index 0.80 (0.08) a 0.74 (0.06) b 0.78 (0.07) a
Fagus sylvatica 0.84 (0.08) a 0.80 (0.06) a 0.85 (0.09) a
Other species 0.72 (0.19) a 0.68 (0.20) a 0.76 (0.15) a
Gini coefficient 0.27 (0.09) b 0.24 (0.06) b 0.40 (0.11) a
Fagus sylvatica 0.25 (0.09) ab 0.23 (0.05) b 0.33 (0.08) a
Other species 0.17 (0.07) b 0.15 (0.09) b 0.22 (0.12) a
Shape index 0.49 (0.22) a 0.49 (0.15) a 0.47 (0.08) a
Fagus sylvatica 0.56 (0.24) a 0.64 (0.12) a 0.57 (0.15) a
Other species 0.80 (0.21) a 0.90 (0.23) a 0.65 (0.22) a
Significant differences among stand types (KruskalWallis non-parametric test, p < 0.05) are indicated by different letters. The values are means and standard
deviations (in brackets).
 A. Merino et al. / Forest Ecology and Management 250 (2007) 206214
Fig. 2. Pooled tree diameter distributions in the three stand types. Data pooled
from all sample plots of each stand type.
(r = 0.39, p < 0.01) and the type of subsurface soil horizon (A2,
AB, B, . . .). The lowest C contents were thus found in the
shallowest and stoniest soils (Leptosols), with less than
60 Mg C ha1. On the contrary, umbrisols, with deep, organic
matter-rich A umbric horizons, showed the largest C stocks, of
up to 250 Mg C ha1 (Fig. 4d). The soil C pool distribution
differed among the three types of soils (probability value for the
KolmogorovSmirnov test, p < 0.05 for any combination). The
design of the study did not allow detection of changes in soil C
storage that were attributable to forest management.
211
Table 3
Results of the KolmogorovSmirnov goodness-of-fit test comparing the dbh-
distributions between different stand types
KolmogorovSmirnov two-sample Ksa Pr > Ksa
test (asymptotic)
Unmanaged mature stands compared 1.78 0.0035
with partially cut stands
Fagus sylvatica 2.41 <0.0001
Unmanaged mature stands compared 2.31 <0.0001
with abandoned pollard stands
Fagus sylvatica 2.19 0.0001
Partially cut stands compared 3.09 <0.0001
with abandoned pollard stands
Fagus sylvatica 2.88 <0.0001
4. Discussion
The mean tree (>7.5 cm dbh) densities observed ranged
from 445 stems ha1 in pollard to 758 stems ha1 in the
partially cut stands. These values are higher than those observed
by Meyer et al. (2003) in three Albanian virgin beech forests
(320388 stems ha1) and by Oheimb et al. (2005) in a near-
natural beech forest in Germany (263 stems ha1). However,
the beech forests in the latter studies consisted almost
exclusively of F. sylvatica (98 and 99% of total stems ha1,
respectively). In addition, the different sampling method and
plot size used makes quantitative comparison between different
studies difficult. When only the number of beech stems per
hectare is taken into account, the differences disappear, except
for PC stands in which a higher density is maintained
(676 beech stems ha1).
The basal area values observed in the present study,
considering only beech trees, are also of the same order of
magnitude as those observed for virgin beech forests or near-
natural beech forests in Central Europe (Meyer et al., 2003;
Oheimb et al., 2005), except for pollard stands in which the
basal area is higher (89.6 m2 ha1) because of the abundance of
old and large trees resulting from the silvicultural treatments
used.
The number of old and large trees is one of the features most
frequently used to characterise generic natural beech forests
(Rademacher et al., 2001). The mean number of old and large
living trees (dbh > 80 cm) in unmanaged stands in the present
study was 14 trees ha1. This value is very close to those
observed by Meyer et al. (2003) in three Albanian virgin beech
forests (519 trees ha1) and those reported for some of the
most important beech-dominated, near-natural stands in France
(1223 trees ha1); Denmark (814 trees ha1) and Germany
(13 trees ha1) (Koop and Hilgen, 1987; Emborg et al., 2000;
Oheimb et al., 2005). Again, the exceptionally high number of
old and large trees observed in the pollard stands
(57 trees ha1) is the result of particular management inter-
ventions.
The diameter distributions in unmanaged stands generally
showed a reverse J-shaped pattern. Similar results have been
found for three virgin forests in Albania (Meyer et al., 2003).
212 A. Merino et al. / Forest Ecology and Management 250 (2007) 206214

Fig. 3. Mean amounts of C (Mg ha1) in biomass and soils for different soil types. (UM) Unmanaged mature stands; (PC) partially cut stands; (AP) abandoned
pollard stands. Since the allometric equations were not suitable for pollard trees, they were not applied for calculating the tree biomass in these stands.

However, a bimodal pattern was observed for some near-natural
forests in Central Europe, with a second maximum ranging
from 100 to 180 years (Emborg et al., 2000). The latter author
suggested that the bell-shaped section of the diameter
distribution at these ages reflects large-scale beech regeneration
due to a natural phase of decline and regeneration (e.g. La Taille
forest in France or Serrahn forest in Germany) or after cattle
grazing ceased (e.g. Suserup Skov forest in Denmark).
The diameter distributions of PC stands showed the highest
number of small recruits and a lack of full-size canopy trees as a
consequence of past cutting and current regeneration. The
selective felling of large trees in these stands led to successful
regeneration because of the creation of large gaps not readily
covered by crown expansion of subdominant trees.
The total C stock in these systems, and the partitioning over
the different compartments, were determined by the past forest
management and the type of soil. The average C pool in above
ground tree biomass in the partially cut forests, 129 Mg ha1,
was comparable to those reported for similar types of forest in
Belgium (123 Mg ha1, Vande Walle et al., 2001), Germany
(156 Mg ha1, Holscher et al., 2001; 120161 Mg C ha1,
Joosten et al., 2004) and higher than in beech forests in other
Spanish regions (66 Mg ha1, Santa Regina and Tarazona,
1999). With respect to other forest systems in the region, the
tree C stocks are similar to those in near-natural Q. robur forests
(123 Mg ha1, Balboa et al., 2006). In the unmanaged stands,
the mean C pool in the aboveground tree biomass was almost
100 Mg ha1 (205 Mg ha1) higher than in the partially cut
stands, which is attributable to the large proportion of trees
larger than 50 cm in these systems. This is consistent with the
idea that the C stocked in tree biomass increases with the stand
age and with the number of larger trees (Vanninen et al., 1996).
The results of the present study suggest that the current
recovery from past moderate human disturbances may lead to
substantial increases in the C storage of live biomass in these
forests, which has also been suggested in tropical forests (Vieira
et al., 2004).
However, it is not known if this trend will undergo an abrupt
change in the near future, as observed in other parts of Europe
(Emborg et al., 2000; Rademacher et al., 2001), as no cyclical
dynamic model has yet been established for Iberian beech.
Application of this type of model would possibly show that the
structural and dasometric characteristics of stands, such as AP
would resemble a degradation stage, whereas those of the other
types of beech stands (PC and UM) would resemble early
biostatic and late biostatic stages, respectively. Specific studies
are therefore required to determine the structural typology and
the regime of non-anthropogenic disturbances that affect these
forests, as well as the relationships with C sequestration in the
long-term.
Despite the effect of altitude on net primary production,
which has already been described in different forest ecosystems
(e.g. Waide et al., 1998), no relationship between altitude and
tree biomass was observed for any of the types of forest
considered. The complexity of this mountainous landscape,
with regards to slope, orientation, soil depth and microclimatic
situation make it difficult to establish relationships of this type.
The increase in altitude is not gradual, since there are valleys

Table 4
Parameters of the Weibull function of the three stand types for pooled tree data and for Fagus sylvatica
Unmanaged (UM) Partially cut (PC) Abandoned pollard (AP)
Scale parameter (b) 4.53 (1.88) b 3.07 (0.67) c 8.61 (3.14) a
Fagus sylvatica 4.77 (1.92) b 3.16 (0.75) c 8.50 (3.07) a
Shape parameter (c) 0.45 (0.03) ab 0.44 (0.01) b 0.49 (0.04) a
Fagus sylvatica 0.45 (0.03) a 0.44 (0.01) a 0.47 (0.03) a
Significant differences among stand types (KruskalWallis non-parametric test, p < 0.05) are indicated by different letters. The values are means and standard
deviations (in brackets).
 A. Merino et al. / Forest Ecology and Management 250 (2007) 206214
Fig. 4. Probability density distribution of the carbon contents in the different C pools considered in this study (total tree biomass, aboveground tree biomass, litter
layer and mineral soil) estimated with a Gaussian kernel-smoothing technique using the direct plug-in method to compute the bandwidth.
within the area. The frequent fog and heavy rains in the valleys
significantly modify the microclimatic conditions, thereby
affecting primary production.
The mass of organic litter material was considerably lower
than reported in other colder areas (e.g. Vande Walle et al.,
2001), reflecting the lower residence time of the litter. The
present results are consistent with the findings of Meier et al.
(2005) in that there was no evidence of smaller amounts of litter
mass in nutrient-poor soils. The lower amount of litter layer in
the partially cut forests was due to the lower litterfall
production as a consequence of the lower aboveground tree
biomass and lower tree age and possibly the accelerated
decomposition rates resulting from the different forest structure
(canopy density and species). Changes in microclimatic
conditions (higher soil humidity and temperature) and the
lower proportion of other broadleaf species may have modified
the rate of litter decomposition in the gaps (Zhang and Zak,
1995; Pardo et al., 1997). However, it has been observed that
increased litter fall from the rapidly re-established vegetative
cover prevents the decline in organic litter layer matter after
cutting (Lee et al., 2002).
5. Conclusions
The diversity in stand structure of the southern European
remnant beech forests under study are comparable to that
213
described in central European near-natural beech forests. The
potential C storage in these systems is very high, especially in
the unmanaged forests, which include large, old live trees. The
recovery from past moderate human disturbances should lead to
substantial increases in the C storage. The value of ecosystem
functions should be taken into account in evaluating forestry
decisions in such fragile near-natural forests. Protective
measures undertaken in old managed stands should enhance
biodiversity and the role of both soil and tree components as
long-term C sinks.
Acknowledgements
The authors thank Mr. Miguel Angel Negral, Manuel
Fontao, Santiago Grandas and Javier Ferreiro for their
assistance in the field work. The English grammar of the text
was revised by Dr. Christine Francis.
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