Open Journal of Forestry
2012. Vol.2, No.4, 200-206
Published Online October 2012 in SciRes (http://www.SciRP.org/journal/ojf) http://dx.doi.org/10.4236/ojf.2012.24024
Copyright © 2012 SciRes.
Biodiversity in Chestnut Woodlots: Management Regimen vs
Javier Guitián1, Pablo Guitián 1, Ignacio Munilla1, José Guitián2, Josefina Garrido3,
Liliana Penín3, Paula Domínguez1, Luis Guitián4
1Department of Botany, University of Santiago, Santiago, Spain
2Department of Celular Biology and Ecology, University of S an ti ago , Santiago, Spain
3Department of Ecology and A nimal Biology , University of Vigo, Vigo, Spain
4Deparment of Geography, University o f S a nt i ag o , Santiago, Spain
Received July 19th, 2012; revised August 20th, 2012; accepted September 1st, 2012
This paper analyzes the effect of woodlot size and land-use intensity on the species richness of vascular
plants, birds, beetles, and ants in Castanea sativa (chestnut) woodlots of the northwestern Iberian Penin-
sula included in the category “9260 Castanea sativa woodland”, “Annex I, DC 92/43/European Commu-
nity”. The results show that the surface area of the woodlot did not affect the richness of vascular plants
and ants but did affect birds and beetles. The level of abandonment of the woodlot affected only the rich-
ness of vascular plants, while the use level had no significant impact on species richness of any of the
groups. The degree of maturity of the woodlot, estimated by the tree-trunk circumference, determined
only the richness of plants but not that of different groups of animals. In conclusion: 1) Plants and animals
responded differently to woodlot size, abandonment, and the degree of maturity of the woodlots; 2) Tradi-
tional agricultural practices do not negatively affect the biodiversity of the chestnut woodlots of the
northwestern Iberian Peninsula or favor plant diversity; and 3) A traditional use of these woodlots may
continue to play an important role in maintaining the diversity of plant species in the area.
Keywords: Vascular Plants; Birds; Beetles; Ants; Species Richness; Castanea Woodlands
In changing landscapes, the spatial distribution of species
depends on their biological characteristics and the pattern of
land alteration. When land abandonment occurs, the difficulties
in predicting the effects of landscape change on biodiversity
arise from large differences in species’ dispersal behavior and
the unpredictability of patterns of land-use change (Veldkamp
& Lambin, 2001 and references therein). The major problems
related to the abandonment of agricultural land are biodiversity
loss, increased fire frequency and intensity, soil erosion and
desertification, loss of cultural and/or aesthetic values, lower
landscape diversity, and reduced water provision. The aban-
donment of agricultural land may also benefit humans, includ-
ing passive revegetation and active reforestation, water regula-
tion, soil recovery, nutrient cycling, and increased biodiversity
and wilderness (see Rey Benayas et al., 2007 and references
The land-abandonment process generates a mosaic of uses
ranging from complete abandonment to different levels of in-
tensity of exploitation, with potential consequences for the
biological richness of these areas. The consistency of species
responses to land-use changes may be altered by within-patch
habitat conditions. Indeed, the degree of natural and anthropo-
genic disturbance at a small scale may influence species dy-
namics to an equal or greater extent in comparison with pertur-
bation in a larger-scale landscape structure (Debuse et al.,
Throughout most of the northwestern Iberian Peninsula, ma-
ture indigenous forests (e.g., Quercus robur L., Fagus sylvatica
L., etc.) have been intensively exploited and are currently very
scarce. A common formation in this region is the chestnut
(Castanea sativa Mill.) woodland, surrounding small villages
and managed for sweet chestnuts and timber. This is the most
important remaining deciduous woodland type in this region,
and for centuries has in many respects acted as a substitute for
the indigenous deciduous forests throughout most of NW Spain.
Chestnut woodlands are typically comprised of large trees,
many well over 200 years old, but in recent decades these for-
ests, being traditionally exploited for feeding humans and ani-
mals, have suffered a progressive process of abandonment as a
result of rural depopulation and fungal diseases. This has seri-
ously damaged the trees and reduced crop quality (Pitte, 1986),
with economic and ecological consequences (Arnaud et al.,
Several authors have studied the effects that this process, and
the resulting management regimes, have on the plant and ani-
mal richness in chestnut woodlands, showing a clear relation-
ship between the two, and proposing strategies for biodiversity
conservation both locally and regionally (Roberts & Gilliam,
1995; Peltzer et al., 2000; Romane et al., 2001; Hansson, 2001;
Gondard et al., 2001, 2006, 2007; Mason & MacDonald, 2002).
Also, several works have analyzed the influence of spatial
characteristics (area, shape, etc.) on the richness of plants and
animals in these woodlands, although the results are not always
conclusive (Dzwonko & Loster, 1992; Bastin & Thomas, 1999;
Petit et al., 2004; Cousins & Aggemyr, 2008; Konstantinidis et
J. GUITIÁN ET AL.
In this study, we focus on four features that influence species
richness in woodland patches and analyze to what extent the
size and the use of chestnut woodlots affect the richness of
vascular plants, birds, ants, and beetles. For this, we used the
surface area of the woodland, the abandonment of the woodland
in the recent past, as reflected by the structure of its vegetation,
and the current use as determined by the intensity and homo-
geneity of chestnut harvesting. In addition, we analyzed to what
degree forest maturity influences species richness of these
groups. Specifically, after characterizing the woodlots and ana-
lyzing species richness of the different groups, we tried to re-
spond to the following questions: 1) Does the surface area of
the woodlot affect species richness of the different groups? 2)
Does the current level of abandonment and use affect the dif-
ferent groups? 3) Does the degree of maturity of the woodlot
affect species richness of different groups? 4) Finally, in the
three cases analyzed, is the response consistent among the plant
and animal groups studied?
This approach is relevant because managed chestnut wood-
lands are included in the Habitats Directive of Spain (Annex 1
of Directive 92/43: “Castanea sativa woodlands 9260”) and
several studies have shown their high ecological importance in
supporting a wide variety of flora and fauna (Rodriguez-Guitián
et al., 2005; González-Varo et al., 2008).
The Study Area
The study was conducted in the O Courel and Ancares
mountains (provinces of Lugo and León, NW Spain), a highly
cultivated mountain area covering roughly 500 km2. The terri-
tory is part of the Natura 2000 Network included in the “An-
cares-Courel Site of Community Importance” (ES1 120001).
The landscape is made up of a mosaic of small villages, farm-
land, and chestnut woodlots set into a matrix of thickets and
woodlands dominated by Quercus, Betula, and Fagus.
Woodland Ch aracterizatio n
Of the approximately 100 woodlots found in the area (un-
published data), we selected 30 woodlots of different sizes and
land-use intensity. The woodlands were spatially characterized
by generating a series of map layers using Geomedia (Inter-
graph) software supported by GVSIG (free software) for areas
outside the Autonomous Community of Galicia. These founda-
tions were used to generate a geographic information system
with UTM projection (zone 29), based on the European 1950
datum and in two formats (shape and mdb) that can be read by
any standard geographic information system. A photo interpre-
tation was performed to define the edges of the chestnut groves,
backed by field inspection and based on the orthophoto SIX-
PAC of the area taken in 2002-2003.
In the woodlots, two different management levels were dis-
tinguished: the level of abandonment and the level of current
use. The level of abandonment was quantified by the direct
observation of the vegetation structure and by interviews with
the local population and woodland owners. On the basis of the
information compiled, three levels were assigned: 1 = aban-
doned or semi-abandoned, the land is not cleared of vegetation
on a yearly basis, but nuts are gathered sporadically; 2 = mod-
erate use, the land is cleared yearly or twice a year; and 3 =
intensive use, the land is cleared of vegetation by burning or
cutting. To verify that the levels of land use assigned to the
woodlands corresponded to their structural characteristics, we
determined the structure of the vegetation in a sample of 16
chestnut woodlots. For this purpose, we located the central
point of each woodlot with the help of an aerial photograph
(Aerial photograph after 2002. SIXPAC, SIGPAC Junta Cas-
tilla y León). The slope was estimated at this point with a cli-
nometer and the slope’s orientation was determined by means
of a compass. Next, four 50-m transects were randomly taken
and, on each transect, five points were randomly assigned to
calculate the vegetative cover index in the strata between 0 and
12m (Karr, 1971). The cover index was calculated as the per-
centage of actual contacts out of all the possible contacts (4
transects × 5 points/transect × 6 strata/point). The cover indices
computed on the basis of the structural data correlated signifi-
cantly with the use levels estimated in the field; hence the latter
indices were used in the a nalyses.
The current use level was quantified on the basis of the pro-
portion of chestnuts with respect to the number of cupules pre-
sent in the woodlot (spiny cupules containing 3 - 7 chestnut
fruits), and the coefficient of variation; this gives an idea of
whether or not the crop was collected. For this, in each woodlot,
random samples were taken from 20 or 12 squares of 2 × 0.5 m
according to the size of the woodlot, counting the number of
chestnuts and cupules present; this ratio was higher in the
woodlots where the collection of chestnut fruits and conse-
quently also its use level proved low. Similarly, the woodlots in
which the current use was intensive should show strong uni-
formity in the degree of intervention, and therefore the coeffi-
cient of variation of the ratio chestnuts/cupules (use heteroge-
neity) was low; in the contrary case, the woodlands having an
intermediate use level should show a higher variation coeffi-
The degree of maturity of the woodlots was estimated on the
basis of the breast-height trunk circumference of the trees. For
this, over the transect, every 25 m, we situated 5 × 5 m squares
alternately on the left and on the right. With a measuring tape,
we measured the circumference of all the trees in each square.
Vascular plants. To determine the number of plant species
in each of the 30 woodlots, we marked a point in the center of
the woodland and drew a circle with a 20-m radius. We identi-
fied and recorded all the vascular plant species found in the
area (total area sampled 3.14 × 202 = 1256 m2). This inventory
was conducted twice a year, in early spring and early summer,
to make sure that all the species found in the plot site were
included in the census. The area selected for the study had been
previously determined by plotting a species area-curve using
5-m radius increments until reaching a value where the number
of species stabilized. The total number of species was then
revised based on the list of indicator species of these woodlands
in the study region (see Rodriguez-Guitián, 2004; Rodríguez-
Guitián et al., 2005, for additional information).
Birds. For the study of the composition and structure of the
bird communities, a sample of 20 woodlots were chosen from
those previously used, and were visited in the summer (nesting)
and winter (overwintering) for 3 years (2006, 2007, and 2008).
A total of 240 censuses were made (20 woodlots × 2 seasons ×
3 years × 2 replicates), counting 2702 individuals of 39 species.
Ants. To study the ants, we used 8 woodlots in which we
Copyright © 2012 SciRes. 201
J. GUITIÁN ET AL.
randomly drew 30-m-long transects the length of the woodlot.
Along these lines, we placed 30 pitfall traps, which consisted of
a plastic bottle c. 10 cm in diameter × 15 cm deep containing
detergent surfactant so the ants would sink in the liquid. After
the traps were left for 24 h, the contents were collected in 70˚
alcohol for preservation and subsequent identification. This
sample was made in 2006 and 2007.
Beetles. In the study of the beetles, eight woodlots were
sampled in an attempt to collect all the variability possible in
the size and degree of use of these woodlots. In each woodlot,
pitfall traps (5 - 10) were set at a given number of random
points set, depending on the surface area, this constituting one
of most widely used indirect techniques for studying Coleop-
tera populations (Ribera et al., 2001; Taboada et al., 2004). The
traps used in this study consisted of a plastic bottle half filled
with a liquid preservative (Propilenglicol at 25%); the bottle
measured 12 cm high and 8 cm in diameter and was buried in
such a way that its mouth was higher than or flush with the soil
level, while a wooden shield protected against falling plant
debris. The contents were collected after approximately two
weeks. Traps were set during summer and autumn of two con-
secutive years (2006 and 2007) at the 61 sampling points pre-
viously established. The trap contents were taken to the labora-
tory, preserved in 70˚ alcohol for subsequent identification and
Species richness was assessed by means of individual-based
rarefaction (Gotelli & Colwell, 2001), thus accounting for dif-
ferences in sampling effort. Datasets were standardized to a
common number of individuals (30 for birds and ants and 100
for beetles) assuming a hypergeometric sampling distribution
(Heck et al., 1975).
The effects of the different variables on the total number of
species were analyzed by General Linear Modeling (GLM,
probability distribution Poisson, link function Logarithm). As
dependent variable, the number of species of each group was
used, calculating the mean for the biennial samplings. The level
of abandonment was included as the factor and the rest of the
predictor variables were included in the model as covariables.
All of the analyses were carried out with the statistical pack
Characteristics of the Woodlands
Chestnut woodlots are found at altitudes ranging from 600 to
1000 s nm. The area of the woodlots ranged from 0.3 to 94
hectares. Some 53% of the woodlots had a low level of aban-
donment, the mean being 20%, with 27% being abandoned. The
level of current use estimated by the relationship of chest-
nuts/cupules varied from 0.43 for the highest use level and 2.20
for the lowest; the CV (mean/sd × 100) of this ratio varied from
147.43 for the most heterogeneous forest to the use level of
16.36 for the most homogeneous. The circumference of the
trees ranged from 36.76 ± 52.47 cm in the youngest woodlot to
329.62 ± 110.37 cm in the oldest woodlot.
Species Richness of Different Taxa
The number of species of plants, birds, ants and beetles in the
sweet chestnut woodlots are show in Table 1.
Vascular plants. In the woodlots studied, we identified in
total nearly 160 species of vascular plants. The total number of
species in each woodlot ranged from 6 to 32. In addition to the
chestnut, the most frequent woody species were Fraxinus ex-
celsior and Quercus robur, present in more than 75% of the
woodlands sampled, with Luzula campestris and Viola riviniana
being the most common understory plants. The complete list of
species appears in Guiti a n e t al. (2012).
Birds. A total of 39 species and 2702 individuals were re-
corded. The most common species were robin (Erithacus
rubecula), wren (Troglodytes troglodytes), and blackbird
(Turdus merula) with, respectively, 14.9, 12.0, and 11.5% of
sightings. The (accumulated) total number of species per
woodlot ranged from 9 to 20. When rarefied to a common
number of individuals, mean species richness was 11.3 (6 - 14).
(See Appendix 1 for the complete list of species)
Beetles. A total of 3.582 individuals were identified, be-
longing to 23 families and 78 species. Carabidae (1152 speci-
mens), Anobiidae (974), and Staphylinidae (670) were the
families with the largest number of specimens. Taking into
account the number of species in each family, Staphylinidae (13
species), Carabidae (12), Curculionidae (9), and Leiodidae (7)
were noteworthy. The results show that Ptinus fur (Anobiidae)
was the most abundant species captured (926 specimens). The
total number of species in each woodlot ranged from 10 to 28.
(See Appendix 1 for the complete list of species).
Ants. The ant samplings resulted in the identification of a
total of 4078 individuals in the year 2006 and 1362 in 2007.
The individuals captured corresponded to the genera Formica,
Lasius, Myrmica, Tapinoma, Aphaenogaster, and Leptothorax.
The most abundant species was Formica rufa. The total number
of species in each woodlot ranged from 1 to 7 (See Appendix 1
for the complete list of species).
Relation between Species Richness with Surface Area
of the Woodlots
The analysis showed that the surface area of the woodlot had
no significant effect on vascular plant richness (Table 2), but
did have an impact on the richness of birds and beetles; contrar-
ily, no effect was found on ants (Table 2). In the first two cases,
the effect was positive, so that the largest woodlands were
richer in birds and beetles (Figure 1).
Relationship of Species Richness with the Level of
Abandonment and the Level of Current Use
Species richness of vascular plants apparently responded to
the level of abandonment but not with the level of current use,
both considering the quotient chestnuts/cupules as well as its
Number of species of plants, birds, ants and beetles in the sweet
chestnut woodlots of Caurel. N = number of woodlots examined. SD =
N TotalMinimum Maximun MeanSD
Plants30156 6 32 19.2 6.1
Birds 2039 9 20 10.9 4.1
Ants 8 10 1 7 4.3 2.2
Beetles8 78 10 28 23.4 6.2
Copyright © 2012 SciRes.
J. GUITIÁN ET AL.
coefficient of variation (use heterogeneity; Table 2). As a
whole, the GLM model proved to be significant (p < .05) and
the results show that the level of land use had a significant ef-
fect on total plant species richness, with a greater richness
found in the moderate land-use level (mean 13.8, 20.5 and 19.7
for levels 1, 2, and 3 re spectively).
Species richness for birds, ants, and beetles was not deter-
mined by the level of abandonment nor the level of current use
estimated by the chestnuts/cupule ratio as their coefficient of
variation (Table 2).
Relationship between Species Richness and Forest
Only the richness of the vascular plants showed a significant
relation with woodlot maturity, in such a way that the most
mature woodlot had greater species richness. This variable had
no significant effect on the richness of any of the animal groups
studied (see Table 2).
Results of GLM to analyze the effects of the different variables on the
total number of plant and animal species.
Wald Chi-square Df p
(Intersection) 164.846 1 .00
Abandonment 8.742 2 .01
Current use .525 1 .47
Area .668 1 .41
Age 3.574 1 .05
Current use (CV) .094 1 .76
(Intersection) 32.381 1 .00
Abandonment .096 2 .95
Current use 2.148 1 .14
Area 15.406 1 .00
Age .853 1 .36
Current use (CV) .097 1 .76
(Intersection) 4.610 1 .03
Abandonment .286 2 .87
Current use 2.821 1 .09
Area 3.742 1 .05
Age .755 1 .39
Current use (CV) .096 1 .76
(Intersection) 3.256 1 .07
Abandonment .025 2 .99
Current use .311 1 .58
Area 2.586 1 .11
Age .004 1 .95
Current use (CV) .429 1 .51
Relationship between woodlot size (hectares) and species rich-
ness of different groups: (a) Vascular plants; (b) Birds; (c) Bee-
Castanea sativa woodlands in the northwestern Iberian Pen-
insula vary widely in surface area and use intensity, which may
Copyright © 2012 SciRes. 203
J. GUITIÁN ET AL.
determine both the number and abundance of plant and animal
species. This variation does not appear to be related to differ-
ences in local environmental conditions, since the woodlots are
generally located on deposits in areas of Quercus pyrenaica
woodlands in the mountains of the western Iberian Peninsula.
In general, our results revealed major differences in species
richness between woodlots, and showed that in traditionally
managed woodlots, vascular plants, birds, beetles and ants,
were likely to respond differently to forest-patch size, age, and
Relation of Species Richness to the Surface Area and
the Use Level of the Woodlots
Our results show that the surface area of the woodlots bore
no relation to species richness of the vascular plants, in agree-
ment with the findings of Guitián et al. (2012), but was deter-
minant in the case of birds and beetles though not for ants. In
the same geographical setting, Gonzalez Varó et al. (2008)
found that woodland size was the only variable that signifi-
cantly predicted the presence of breeding pairs de Sitta eu-
ropaea in these forests, and that the number of pairs was
strongly predicted by woodlot size; in the same sense, Baileya
et al. (2002) reported evidence for patch area and composition
effects: larger woodlands support more woodland bird species.
In beetles, responses have been show in several ways. Gibb
and Hochuli (2002) demonstrated that the large fragments had
no more species per unit of area in most of the arthropod groups,
while Fujita et al. (2008) found a decline in the number of
Carabidae and Brachinidae species in small fragments of forests
in Japan. In the case of ants, Gibb and Hochuli (2002) indicated
that small fragments were richer in species.
Relation of Species Richness with the Level of
Abandonment and Current Use
Our results reflect that the current use did not affect species
richness in any of the animal groups studied, but it does affect
plants. In general, the works that have analyzed this issue
agreed with the intermediate disturbance hypothesis: moderate
management would safeguard most potential plant species
(Hansson, 2001; Dollman et al., 2007). Gondard et al. (2007)
reported similar results in two chestnut woodlands of the Medi-
terranean region and proposed the maintenance of a mosaic of
human-altered woodlands to maintain regional biodiversity. In
the chestnut woodlands of England, Mason and McDonal (2002)
found greater species richness and a denser vegetative cover
two or three years after clearing work was performed, with
declining numbers in subsequent years.
On the contrary, the results failed to show an effect by the
level of abandonment and current use in the richness of the
animal groups studied. In studies concerning the joint effect of
the landscape structure and land use on animal richness, group
responses were found to vary. Thus, Atauri and de Lucio (2001)
showed that the response of species richness to landscape het-
erogeneity varies depending on the group of species considered.
For birds and Lepidoptera, the most important factor affecting
the distribution of species richness is landscape heterogeneity,
while other factors, such as the speciﬁc composition of land use,
prove secondary at this scale. On the other hand, amphibian and
reptile richness is more closely related to the abundance of
certain land-use types. Bailey et al. (2002) correlated bird and
mammal distributions to landscape scale measures of fragment
distribution, in contrast to butterfly and plant species, which
exhibited a stronger correlation to patch- scale measures.
For ants, there is evidence that the internal characteristics of
the forest (use) can have more influence than its size or the
spatial configuration of the landscape. Thus, for example, De-
buse et al. (2007), in a study on ant richness in Australian for-
ests, found that within-patch habitat characteristics explained
more than twice the amount of ant species variation attributable
to fragmentation and four times the variation explained by
A reasonable explanation could stem from the fact that we
are dealing with woodlots that have remained stable over the
course of the last three centuries (“seminatural ancient wood-
lands”) in a setting where traditional agricultural practices are
being abandoned. This could help them maintain a high bio-
logical richness, regardless of their size and degree of connec-
tion with other forest masses (Guitián et al., 2012).
Relation of Species Richness to Forest Maturity
In the study region, the age of chestnut woodlots proved de-
terminant for vascular-plant richness. Previous studies have
reported similar responses and, in some cases, contrary, de-
pending on the type of forest considered (Jacquemin et al., 2001;
Widenfalk & Weslien, 2009 and references therein). Many
authors have shown that historical land use may have long-term
effects on present-day environmental conditions of forests,
hampering or slowing down the colonization processes of many
forest plant species. Forest age also represents the colonization
period which is directly related to the probability of a species
reaching a forest fragment (see references in Jacquemin et al.,
From a different perspective, bird, beetle, and ant richness
can not be determined by the degree of forest maturity appar-
ently because the degree of forest maturity may determine spe-
cies composition and abundance of the different groups more
than the number species (se e , e.g. Vasconcelos, 1999).
In the case of birds, Ficetola et al. (2006) found that in forest
fragments the vertebrate species responded in different ways to
perturbations and that, of the bird species studied, only the
presence of Parus caeruleus was related to forest maturity. The
use of carabids to estimate woodland age was investigated in
deciduous woodlands in the UK for a seven-year period (Ter-
rell-Nield, 1999). A significant relationship was found between
the age of woodland and the richness of the carabid fauna, but
not between the woodland area or circumference and carabid
diversity. However, it is important to consider that different
results were found according to the food habitats of the differ-
ent beetle groups (saproxylic vs non-saproxylic).
In conclusion, our results show that vascular plants and ani-
mal groups studied respond differently to woodlot size, aban-
donment level, and woodlot use. This provides substantial evi-
dence that traditional practices do not negatively affect the
biodiversity of the chestnut woodlots of the northwestern Ibe-
rian Peninsula or favor plant diversity. Therefore, a traditional
use of these woodlots may continue to play an important role in
maintaining the diversity of plant species in the area.
We would like to express our gratitude to Tania Veiga, Car-
men Docampo, Ainhoa Magrach, Alberto Tinaut and Asier R.
Larrinaga for their help at various stages of the project. The
Copyright © 2012 SciRes.
J. GUITIÁN ET AL.
study of the Chestnut Woodlands was funded by the “Dirección
Xeral de Investigación of the Xunta de Galicia” through the
sectorial research program on Biodiversity. The “Dirección
Xeral de Medio Rural” provided the human and material re-
sources for the execution of the cartography of the woodlands.
Arnaud, M. Y., Chassany, J. P., Dejean, R., Ribart, J., & Queno, L.
(1997). Economic and ecological consequences of the disappearance
of traditional practices related to chestnut groves. Journal of Envi-
ronmental Management, 49, 373- 391. doi:10.1006/jema.1995.0120
Atauri, J. A., & de Lucio, J. V. (2001). The role of landscape structure
in species richness distribution of birds, amphibians, reptiles and
lepidopterans in Mediterranean landscapes. Landscape Ecology, 16,
Baileya, S.-A., Haines-Young, R. H., & Watkins, C. (2002) Species
presence in fragmented landscapes: Modeling of species require-
ments at the national level. Biolog ical Conservation, 108, 307-316.
Bastin, L., & Thomas, C. D. (1999). The distribution of plant species in
urban vegetation fr a gments. Landscape E cology, 14, 493-507.
Cousins, S. A., & Aggemyr, E. (2008). The influence of field shape,
area and surrounding landscape on plant species richness in grazed
ex-fields. Biological Conservation, 141, 126-135.
Debuse, V. J., King, J., & House, A. P. N. (2007). Effect of fragmenta-
tion, habitat loss and within-patch habitat characteristics on ant as-
semblages in semi-arid woodlands of eastern Australia. Landscape
Ecology, 22, 731-745. doi:10.1007/s10980-006-9068-0
Dolman, P. M., Hinsley, S. A., Bellamy, P. E., & Watts, K. (2007).
Woodland birds in patchy landscapes: The evidence base for strate-
gic networks. International Journal of Avian Science, 149, 146-160.
Dzwonko, Z., & Loster, S. (1992). Species richness and seed dispersal
to secondary woods in Souther Poland. Journal of Biogeography, 19,
Ficetola, G. F., Sacchi, R., Scali, S., Gentilli, A., De Bernardi, F., &
Galeotti, P. (2007) Vertebrates respond differently to human distur-
bance: Implications for the use of a focal species approach. Acta
Oecologica, 31, 109-118. doi:10.1016/j.actao.2006.10.001
Fujita, A., Maeto, K., Kagawa, Y., & Ito, N. (2008). Effects of forest
fragmentation on species richness and composition of ground beetles
(Coleoptera: Carabidae and Brachinidae) in urban landscapes. Ento-
mological Science, 11, 39- 48. doi:10.1111/j.1479-8298.2007.00243.x
Gibb, H., & Hochuli, D. F. (2002). Effects of forest fragmentation on
species richness and composition of ground beetles (Coleoptera:
Carabidae and Brachinidae) in urban landscapes. Biological Conser-
vation, 106, 91-100. doi:10.1016/S0006-3207(01)00232-4
Gondard, H., Romane, F., Grandjanny, M., Li, J. Q., & Aronson, J.
(2001). Plant species diversity changes in abandoned chestnut (Cas-
tanea sativa) groves in southern France. Biodiversity and Conserva-
tion, 10, 189-207. doi:10.1023/A:1008997625523
Gondard, H., Romane, F., Santa-Regina, I., & Leonardi, S. (2006).
Forest management and plant species diversity in chestnut stands of
three Mediterranean areas. Biodiversity and Conservation, 4, 1129-
Gondard, H., Santa-Regina, I., Salazar, S., Peix, A., & Romane, F.
(2007). Effect of forest management on plant species diversity in
Castanea sativa stands in Salamanca (Spain) and the Cévennes
(France). Science and Research Essays, 2, 62- 70.
González-Varo, J. P., López-Bao, J. V., & Guitián, J. (2008). Presence
and abundance of the Eurasian nuthatch Sitta europaea in relation to
the size, isolation and the intensity of management of chestnut
woodlands in the NW Iberian Peninsula. Landscape Ecology, 23, 79-
Gotelli, N. J., & Colwell, R. K. (2001). Quantifying biodiversity: Pro-
cedures and pitfalls in the measurement and comparison of species
richness. Ecology Letters, 4, 379-391.
Guitián, J., Guitián, P., Magrach, A., Docampo, C., Domínguez, P., &
Guitián, L. (2012). Effect of management and spatial characteristics
on plant species richness of Castanea sativa Mill. Woodlots in the
NW Iberian Peninsula. Journal of Forest Research, 17, 98-104.
Hannson, L. (2001). Traditional management of forests: Plant and bird
community responses to alternative restoration of oak-hazel wood-
land in Sweden. Biodiversity and Conservation, 10, 1865-1873.
Heck, K. L., Van Belle, G., & Simberloff, D. (1975). Explicit calcula-
tion of rarefaction diversity measurement and the determination of
sufficient sample size. Ecology, 56, 1459-1461. doi:10.2307/1934716
Jacquemin, H., Butaye J., & Hermy, M. (2001). Forest plant species
richness in small, fragmented mixed deciduous forest patches: The
role of area, time and dispersal limitations. Journal of Biogeography,
28, 801-812. doi:10.1046/j.1365-2699.2001.00590.x
Karr, J. R. (1971). Structure of avian communities in selected Panama
and Illinois habitats. Eco lo gi cal M on og ra phs, 41, 207-2 33.
Konstantinidis, P., Tsiourlis, G., Xofis, P., & Buckley, G. P. (2008)
Taxonomy and ecology of Castanea sativa Mill. Forest in Greece.
Plant Ecology, 195, 235-256. doi:10.1007/s11258-007-9323-8
Mason, C., & MacDonald, S. M. (2002). Responses of ground flora to
coppice management in English woodland—A study using perma-
nent quadrants. Biodiversity and Conservation, 11, 1773-1789.
Peltzer, D. A., Bast, M. L., Wilson, S. D., & Gerry, A. K. (2000). Plant
diversity and tree responses following contrasting disturbances in
boreal forest. Forest Ecology and Management, 127, 191-203.
Petit, S., Griffiths, L., Samrt, S. S., Stuart, R. C., & Whight, S. M.
(2004). Effect the area and isolation of woodland patches on her-
baceous plant species richness across Great Britain. Landscape
Ecology, 19, 463-471. doi:10.1023/B:LAND.0000036141.30359.53
Pitte, J. R. (1986). Terres de castanide. Hommes et paysages du
châtanier de l’Antiquité à nous jours. Paris: Fayard.
Rey Benayas, J. M., Martins, A., Nicolau, J. M., & Schulz, J. J. (2007).
Abandonment of agricultural land: An overview of drivers and con-
sequences. CAB Reviews, 2, 1-14. doi:10.1079/PAVSNNR20072057
Ribera, I., Dolèdec, S., Downie, I. S., & Foster, G.N. (2001). Effect of
land disturbance and stress on species traits of ground beetle assem-
blages. Ecology, 82, 1112-1129.
Roberts, M. R., & Gilliam, F. S. (1995). Patterns and mechanisms of
plant diversity in forested ecosystems: Implications for forest man-
agement. Ecological Applications, 5, 969-977. doi:10.2307/2269348
Rodríguez-Guitián, M. (2004). Aplicación de criterios botánicos para a
proposta de modelos de xestión sustentable das masas arborizadas
autóctonas de subsector galaico asturiano septentrional. Doctoral
Thesis, Santiago: University of Santiago.
Rodríguez-Guitián, M., Rigueiro, A., Real, C., Blanco, J., & Ferreiro da
Costa, J. (2005). El hábitat “9269 Bosques de Castanea sativa” en el
extremo noroccidental ibérico: Primeros datos sobre la variabilidad
florística de los “soutos”. Bulletin de la Societé d’Historie Naturelle
de Toulouse, 1441, 75-81.
Romane, F., Gondard, H., Grandjanny, M., Grossmann, A., Renaux, A.,
& Shater, Z. (2001) Measuring and managing plant species diversity
in the chestnut (Castanea sativa Mill.) ecosystems of the Cevennes.
Forest, Snow and Land s ape Research, 76, 493-497.
Taboada, A., Kotze, D. J., & Salgado, J. M. (2004). Carabid beetle
occurrence at the edges of oak and beech forests in NW Spain.
European Journal of Entomology, 101, 555-563.
Terrell-Nield, C. (1999). Is it possible to age woodlands on the basis of
their carabid beetle div ersity? Journal of Entomology, 109, 136-145.
Vasconcelos, H. L. (1999). Effects of forest disturbance on the structure
of ground-foraging ant communities in central Amazonia. Biodiver-
sity and Conservation, 8, 409-420. doi:10.1023/A:1008891710230
Veldkamp, A., & Lambin, E. F. (2001). Predicting land-use change.
Agriculture, Ecosystems and Environm e nt, 85, 1-6.
Copyright © 2012 SciRes. 205
J. GUITIÁN ET AL.
Copyright © 2012 SciRes.
Widenfalk, O., & Weslien, J. (2009). Forest ecology and management
plant species richness in managed boreal forests—Effects of stand
succession and thinning. Forest Ecology and Management, 257,
Anthous (sp1, sp2, sp3)
Calathus (Neocalathus) rotundicollis
Carabus oreocarabus amplipennis
Chrysocarabus lateralis lateralis
Cychrus spinicollis spinicollis
Dienerella (Cartoderema) clathrata
Microscydmus (Minimus) minimus
Ocypus (Ocypus) olens
Ptomophagus (P.) tenuicornis