Biodiversity in Chestnut Woodlots: Management Regimen vs Woodlot Size

doi:10.4236/ojf.2012.24024

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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

200

Biodiversity in Chestnut Woodlots: Management Regimen vs

Woodlot Size

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

Email: [email protected]

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

Introduction

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

therein).

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.,

2007).

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.,

1997).

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

al, 2008).

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).

Methods

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-

cient.

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.

Species Richness

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

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

study.

Data Analysis

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

SPSS v.19.

Results

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

Table 1.

Number of species of plants, birds, ants and beetles in the sweet

chestnut woodlots of Caurel. N = number of woodlots examined. SD =

Standard Deviation.

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

202

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

Maturity

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).

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

Plant species

(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

Birds

(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

Beetles

(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

Ants

(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

(a)

(b)

(c)

(d)

Figure 1.

Relationship between woodlot size (hectares) and species rich-

ness of different groups: (a) Vascular plants; (b) Birds; (c) Bee-

tles; (d)Ants.

Discussion

Castanea sativa woodlands in the northwestern Iberian Pen-

insula vary widely in surface area and use intensity, which may

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

abandonment.

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

habitat loss.

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.,

2001).

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.

Acknowledgements

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

204

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.

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Appendix I

Bird Species

Aegithalos caudatus

Anthus trivialis

Buteo buteo

Carduelis chloris

Certhia brachydactyla

Columba palumbus

Corvus corone

Cuculus canorus

Dendrocopos major

Emberiza cia

Erithacus rubecula

Fringilla coelebs

Garrulus glandarius

Lanius collurio

Motacilla alba

Muscicapa striata

Oriolus oriolus

Parus ater

Parus caeruleus

Parus cristatus

Parus major

Phoenicurus phoenicurus

Phylloscopus bonelli

Phylloscopus ibericus

Pica pica

Picus viridis

Prunella modularis

Pyrrhula pyrrhula

Regulus ignicapillus

Serinus serinus

Sitta europaea

Streptopelia turtur

Sturnus unicolor

Sylvia atricapilla

Troglodytes troglodytes

Turdus merula

Turdus philomelos

Turdus viscivorus

Ant species

Aphaenogaster sp.

Formica fusca

Formica lugubris

Formica rufa

Lasius brunneus

Lasius flavus

Lasius fuliginosus

Leptothorax sp

Myrmica ruginodis

Tapinoma erraticum

Beetle Species

Acalles sp.

Agathidium escorialensis

Amthobium sp.

Anchonidium unguiculare

Anthous (sp1, sp2, sp3)

Aphodius fossor

Archicarabus steuartii

Blaps sp.

Bolitobius castaneus

Brachinus crepitans

Brachyderes lusitanicus

Bryaxis sp.

Caenopsis waltoni

Calathus (Neocalathus) rotundicollis

Carabus oreocarabus amplipennis

Catops grandicollis

Chasmatopterus sp.

Choleva sp.

Chrysocarabus lateralis lateralis

Criptophagus puncipennis

Criptophagus setulosus

Cychrus spinicollis spinicollis

Dactylosternum abdominale

Dacne bipustulata

Dermestes haemorrhoidales

Dienerella (Cartoderema) clathrata

Drusilla canaliculata

Falagriusa thoracica

Galeruca tanaceti

Geotrupes stercorarius

Hylis olexai

Leistus oopterus

Lordithon exoletus

Mastigus prolongatus

Megasternun concinnum

Mesocarabus macrocephalus

Microplepus staphylinoides

Microscydmus (Minimus) minimus

Mycetoporus rufescens

Necrophorus vespilloides

Nicrophorus sepultor

Niptus sp.

Nosodendron sp.

Nothiophilus quadripunctatus

Ocypus (Ocypus) olens

Ocypus pecipennis

Onthophagus taurus

Ontophylus punctatus

Orchesia sp.

Orobitis cyanea

Othius punctulatus

Pachytychius sparsutus

Paleonthophagus vacca

Parabolitobius inclinans

Paralister stercorarius

Phosphuga atrata

Platydracus stercorarius

Pselaphus heisei

Pterostichus cantabricus

Ptinus fur

Ptomophagus (P.) tenuicornis

tenuicornis

Quedius latinus

Sciodrepoides watsoni

Silpha sp.

Speonemadus vandalitiae

Steropus gallega

Strophosoma erinaceus

Strophosoma sp.

Strophosoma umbilicatum

Trechus optusus

Triodonta sp.