La ecología trófica de la lechuza común en los ecosistemas agrícolas de Grecia Central: su aplicación a la distribución y abundancia de sus presas

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Colecciones : TD. Ciencias experimentalesDBAPEEQA. Tesis del Departamento de Biología Animal, Parasitología, Ecología, Edafología y Química Agrícola
Fecha de publicación : 2009
Esta Tesis Doctoral tuvo cuatro objetivos principales. Estudiar la ecología trófica de la Lechuza común en los agro-ecosistemas de Grecia central, comparar los resultados con otras dietas Europeas y Griegas, y desde el espectro de la dieta buscar patrones de distribución, abundancia y estructura de las poblaciones de los micromamíferos en la región de Tesalia, y estudiar su uso espacial estacional en respecto al cambio estacional del hábitat agrícola. 31 sitios de nidificación naturales fueron localizados en la llanura de Tesalia. Un total de 124 muestras han sido realizadas y 10.065 egagrópilas fueron coleccionadas, reflejando 2 épocas reproductoras y 2 no reproductoras. Después el análisis de las egagrópilas un total de 29.061 presas fueron identificadas, en su mayoría micromamíferos. 28 variables ambientales fueron cuantificadas que fueron incluidas en el ratio de 2 km alrededor de cada sitio de muestra. El análisis de datos ecológicos multivariante ha sido aplicada utilizando el programa CANOCO y con el uso de Modelos Lineales Generalizables, y el Análisis de Gradientes Directa ha sido aplicado utilizando el Análisis de Redundancias. La Lechuza común en Tesalia ha capturado principalmente topillos, y especialmente durante la época reproductora, demostrando también una selección de hábitat fuerte hacia pequeñas parcelas de tierra que sostenían topillos. Alternativamente durante las épocas no reproductoras ha capturado ratones y ratas forrajeando a otros tipos de hábitat. La depredación sobre las ratas durante la época no reproductora ha sido una de las más altas en Europa Mediterránea. Nuevos datos de distribución se han presentado por todas las 15 especies de micromamíferos, que anteriormente se consideraban como ausentes de la llanura de Tesalia. Crocidura suaveolens y Microtus thomasi han sido correlacionadas con tipos del suelo y textura del suelo, y han sido fuertemente afectadas del cambio estacional del hábitat agrícola. Las especies del genero Mus y Apodemus siendo competidores débiles y más generalistas han diferenciado sus nichos, ocupando tipos de hábitat otros de aquellas especies especializadas y territoriales. Microtus guentheri ha sido la especie más abundante en el primer año del estudio pero su población sufrió un decrecimiento fuerte durante los tres años del estudio, mientras tanto los demás grupos de micromamíferos han aumentado.The PhD Thesis had four main objectives. Study Barn owl s trophic ecology in the agroecosystems of central Greece, compare it with Greek and European diets, and through that diet spectrum search for patterns of distribution, abundance and structure of small mammal populations in the region of Thessaly, and study their seasonal space use in relation to the change of agricultural habitat. A total of 31 Barn owl breeding sites were located in the lowlands of Thessaly. A total of 124 samplings were realized and 10.065 pellets were collected, reflecting concretely two breeding and two non-breeding periods. After pellet analysis a total of 29.061 prey items were indentified, mainly comprised of small mammals. 28 environmental variables were quantified, as they were included in a 2 km ratio around each sampling site. Multivariate analysis of ecological data was applied with the software CANOCO and the use of Generalized Linear Models, and Direct Gradient Analysis was applied using Redundancy Analysis. Barn owl in Thessaly preyed heavily on voles, especially during breeding seasons and it also presented a strong habitat selection towards small habitat patches that sustained voles. Alternatively during non breeding seasons it preyed on mice and rats foraging over different habitat types. Rat predation during non breeding season was among the highest in Mediterranean Europe. New distribution data were presented for a total of 15 small mammal species, previously considered absent from Thessaly plains. Crocidura suaveolens and Microtus thomasi were correlated with soil types and soil texture, and were strongly affected from the seasonal change of agricultural habitat. Mus and Apodemus species as weak competitors and generalists differentiated their niches by occupying habitat types others than those of specialized and territorial species. Microtus guentheri was the most abundant in the first year but its population crashed during the three year study, while populations of all other small mammal groups increased.
Publicado el : domingo, 29 de julio de 2012
Lectura(s) : 107
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UNIVERSIDAD DE SALAMANCA



FACULTAD DE BIOLOGÍA

Departamento de Biología Animal, Ecología, Parasitología, Edafología y Química
Agrícola
(Área de Zoología)



LA ECOLOGÍA TRÓFICA DE LA LECHUZA COMÚN EN LOS
ECOSISTEMAS AGRÍCOLAS DE GRECIA CENTRAL: SU
APLICACIÓN A LA DISTRIBUCIÓN Y ABUNDANCIA DE SUS
PRESAS





TESIS DOCTORAL
SALAMANCA 2009

VASILEIOS ANASTASIOU BONTZORLOS

UNIVERSIDAD DE SALAMANCA



FACULTAD DE BIOLOGÍA


Departamento de Biología Animal, Ecología, Parasitología, Edafología y Química
Agrícola
(Área de Zoología)





LA ECOLOGÍA TRÓFICA DE LA LECHUZA COMÚN EN LOS
ECOSISTEMAS AGRÍCOLAS DE GRECIA CENTRAL: SU
APLICACIÓN A LA DISTRIBUCIÓN Y ABUNDANCIA DE SUS
PRESAS



THE TROPHIC ECOLOGY OF BARN OWL IN THE
AGRICULTURAL ECOSYSTEMS OF CENTRAL GREECE: ITS
APPLICATION IN THE DISTRIBUTION AND ABUNDANCE OF
ITS PREY












TESIS DOCTORAL
SALAMANCA 2009

VASILEIOS ANASTASIOU BONTZORLOS

El Dr. D. José Salvador Peris Álvarez, profesor titular del Departamento de
Biología Animal, Parasitología, Ecología, Edafología y Química Agrícola de la
Universidad de Salamanca, y director de la tesis con título “La ecología trófica de la
Lechuza común en los ecosistemas agrícolas de Grecia central: Su aplicación a la
distribución y abundancia de sus presas”, elaborada por Vasileios Anastasiou
Bontzorlos en el Área de Zoología de la Universidad de Salamanca para optar al
grado de doctor, considera que dicha tesis presenta los requisitos necesarios para ser
defendida públicamente, por lo que:

Autoriza su presentación ante el tribunal correspondiente.




En Salamanca a ……………… de …………….. de 2008











Fdo.: José Salvador Peris Álvarez
(El Director)







Memoria presentada por el Licenciado
Vasileios A. Bontzorlos para aspirar al
Grado de Doctor por la Universidad de Salamanca.









Fdo.: Vasileios A. Bontzorlos
(El Doctorando)














Αφι ερωμ ένο στου ς γον ε ίς μου κα ι στ η ν αδ ε ρ φή μου

Dedicado a mis padres y a mí hermana

Dedicated to my parents and to my sister














i

Contents
Page
Acknowledgements……………………………………………………………………………....... vi
List of Tables ……………………………………………………………………………………... viii
List of Figures …………………………………………………………………………………….. xi

The Barn owl and the Small Mammal Fauna of Greece
Chapter 1, Introduction, Part I …………………………………………………………………… 1
1.1 Objectives and Structure of this Thesis ……… 2
1.2 Barn owl ………………………………………………………………………………………. 3
1.2.1 General Description and Biology ……… 3
1.2.2 World Distribution, Population Trend and Population Status ………………………….. 6
1.2.3 Threats in Europe and Greece …………………………………………………………... 8
1.2.4 The Barn owl in Greece ………………………………………………………………… 12
1.2.5 The Barn owl Population in Thessaly, Central Greece …………………………………. 12
1.3 Small Mammals ………………………………………………………………………………. 13
1.3.1 Small Mammals in the Mediterranean Basin and the Particularities of the Balkan
15
Peninsula ………………………………………………………………………………...
1.3.2 The Status of Non-Flying Small Mammal Fauna in Greece …………………………… 17
1.3.2.1 Order: Erinaceomorpha …………………………………………………………… 18
1.3.2.1.1 Family: Erinaceidae ………………………………………………………….. 18
1.3.2.2 Order: Soricomorpha ……………………………………………………………… 19
1.3.2.2.1 Family: Soricidae ……………. 19
1.3.2.2.2 Family: Talpidae …………….. 21
1.3.2.3 Order: Rodentia …………………………………………………………………… 21
1.3.2.3.1 Family: Sciuridae ……………. 21
1.3.2.3.2 Family: Gliridae ……………………………………………………………… 22
1.3.2.3.3 Family: Spalacidae …………………………………………………………… 23
1.3.2.3.4 Family: Muridae ……………………………………………………………... 23
1.3.2.3.5 Family: Cricetidae …………………………………………………………… 27
1.3.3 Small Mammals and the Agroecosystems of Thessaly ………………………………… 29
1.4 Resumen ……………………………………………………………………………………… 30
Study Area
Chapter 1, Introduction, Part II .…………………………………………………………………... 35
1.5 Study Area …………………………………… 36
1.5.1 Climate ………………………………………………………………………………….. 38
1.5.1.1 Temperature ………………………. 39
1.5.1.2 Precipitation ……………………………………………………………………….. 40
1.5.2 Geology and Soil ………………………. 40
1.5.2.1 Geological Formations …………………………………………………………….. 42
1.5.2.2 Soil Classification ………………… 43
1.5.3 Agricultural Cultivations ……………………………………………………………….. 46
1.5.3.1 Habitat Classification and Land Uses ……………………………………………... 48
1.5.4 Fauna ……………………………………………………………………………………. 53
1.5.4.1 Amphibian and Reptiles …………………………………………………………... 53
1.5.4.2 Birds ……………………………………………………………………………….. 54
1.5.4.2.1 Birds of Prey …………………………………………………………………. 56
1.5.4.2.2 Non – Raptors ……………….. 57
1.5.4.3 Mammals …………………………. 58
1.6 Resumen ……………………………………………………………………………………… 60
ii

Page
The Trophic Guild of Barn owl in Greece: Review, Comparisons, Mainland - Insular Trends
and Niche Breadth
Chapter 2 ………………………………………………………………………………………….. 64
2.1 Introduction …………………………………………………………………………………… 65
2.2 Studied Areas …………………………………………………………………………………. 68
2.2.1 Mainland Diet Studies …………………. 68
2.2.1.1 Evros Delta ……………………….. 68
2.2.1.2 Wetlands of Northern Greece ……………………………………………………... 70
2.2.1.3 Parthenio …………………………………………………………………………... 70
2.2.1.4 Potidaia …………………………… 74
2.2.1.5 Thessaly …………………………………………………………………………… 74
2.2.1.6 Avlona ……………………………………………………………………………... 78
2.2.1.7 Dafni & Humettus …………………………………………………………………. 78
2.2.2 Insular Diet Studies ……………………………………………………………………... 79
2.2.2.1 Corfu ………………………………………………………………………………. 79
2.2.2.2 Kos ………………………………79
2.2.2.3 Astipalaia …………………………. 83
2.2.2.4 Evoia ………………………………………………………………………………. 83
2.2.2.5 Antikythera ……………………….. 87
2.2.2.6 Crete. 87
2.3 Methods and Material ………………………………………………………………………… 88
2.4 Results ………………………………………………………………………………………… 91
2.4.1 General Overview and Descriptive Statistics …………………………………………... 91
2.4.2 Mainland Barn owl Diets ……………………………………………………………….. 102
2.4.3 Insular Barn owl Diets ………………………………………………………………….. 104
2.4.4 Niche Breadth, Geographical Trends and Comparisons ………………………………... 105
2.5 Discussion …………………………………………………………………………………….. 109
2.6 Resumen ……………………………………………………………………………………… 120

The Feeding Ecology of Barn owl in Central Greece. Geographical Tendencies and Seasonal
Comparisons
Chapter 3 …………………………………………. 124
3.1 Introduction …………………………………………………………………………………… 125
3.2 Methods and Material ………………………………………………………………………… 127
3.2.1 Field Methodology …………………… 127
3.2.2 Laboratory Methodology ……………………………………………………………….. 132
3.2.3 Qualitative and Quantitative Methodology …………………………………………….. 134
3.2.3.1 Biomass ……………………………………………………………………………. 134
3.2.3.2 Niche Indices ……………………………………………………………………… 135
3.2.4 Statistical Analysis ……………………138
3.3 Results ………………………………………………………………………………………… 143
3.3.1 The Barn owl Feeding Habits in Thessaly. General Overview ………………………… 143
3.3.1.1 Niche Breadth ……………………………………………………………………... 147
3.3.3 Geographical Tendencies ……………………………………………………………….. 149
3.3.4 Seasonal Comparisons ………………………………………………………………….. 165
3.4 Discussion …………………………………………………………………………………….. 176
3.5 Resumen …………………………………………………………………………………….... 189


iii

Page
Distribution and Structure of Small Mammal Populations in the Agricultural Ecosystems of

Thessaly, Central Greece. Associations with Habitat, Soil Types and Land Uses
Chapter 4 ………………………………………………………………………………………….. 195
4.1 Introduction …………………………………………………………………………………… 196
4.2 Methods and Material ………………………………………………………………………… 198
4.2.1 Statistical Analysis …………………… 200
4.3 Results ………………………………………………………………………………………… 204
4.3.1 Effect of Environmental Gradients on Small Mammal Assemblages. Distribution and
213 Gradual Composition Change …………………………………………………………...
4.3.1.1 Bi-colored shrew ………………….. 213
4.3.1.2 Lesser white-toothed shrew ……………………………………………………….. 215
4.3.1.3 Etruscan shrew …………………………………………………………………….. 217
4.3.1.4 Guenther’s vole …………………… 220
4.3.1.5 East European vole ……………….. 222
4.3.1.6 Thomas’s pine vole ………………………………………………………………... 224
4.3.1.7 Gray dwarf hamster. 226
4.3.1.8 Yellow-necked field mouse ……………………………………………………….. 230
4.3.1.9 Western broad-toothed field mouse ……………………………………………….. 230
4.3.1.10 Long-tailed field mouse ………………………………………………………….. 232
4.3.1.11 Brown rat ………………………………………………………………………… 234
4.3.1.12 Black rat …………………………. 236
4.3.1.13 House mouse ……………………………………………………………………... 238
4.3.1.14 Macedonian mouse ………………………………………………………………. 241
4.3.1.15 Hazel dormouse ………………………………………………………………….. 243
4.3.2 Individual Response of Small Mammal Species on Each Environmental Gradient …… 246
4.3.3 Response of Small Mammal Species, Environmental Variables and Factors, on
251
Latitudinal and Longitudinal Gradients …………………………………………………
4.4 Discussion …………………………………………………………………………………….. 259
4.5 Resumen ……………………………………………………………………………………… 284

Small Mammal Populations in Mediterranean Agroecosystems. Seasonal Fluctuations,

Temporal Space-Use Patterns, and Crop-Rotation Effect. The Case of Thessaly, Central
Greece
Chapter 5 …………………………………………. 290
5.1 Introduction …………………………………………………………………………………… 291
5.2 Methods and Material ………………………………………………………………………… 295
5.2.1 Statistical Analysis …………………… 297
5.3 Results ………………………………………………………………………………………… 301
5.3.1 Seasonal Differences in Small Mammal Abundances ………………………………….. 301
5.3.2 Factor Analysis in Seasonal Terms ……………………………………………………... 312
5.3.3 Ordination Analysis in Seasonal Terms .. 318
5.4 Discussion ……………………………………. 345
5.5 Resumen ……………………………………………………………………………………… 377






iv

Page
General Discussion and Implications for Conservation
Chapter 6 ………………………………………………………………………………………….. 383
6.1 Common Agricultural Policy (CAP) and Biodiversity Issues………………………………… 384
st6.2 Intensified Agroecosystems and Overpopulation in Europe at the 21 Century. Facing the
387
challenge……………………………………………………………………………………...........
6.3 Birds in European farmlands and population trends…………………………………………... 389
6.4 Greek Fauna as Part of Mediterranean Europe, Balkan Peninsula and Border with Anatolia... 391
6.5 Raptors Guild and Small Mammal Assemblages in Agroecosystems of Central Greece…….. 393
6.6 Predator – prey relationships in a dynamic agroecosystem. Proposals for conservation,
397
problems and constraints
6.7 Resumen ……………………………………………………………………………………… 404

Conclusions
Chapter 7…………………………………………………………………………………………... 411
7.1 Conclusions…………………………………… 412
7.2 Conclusiones………………………………….. 416

Bibliographical References………………………………………………………………………... 422
Appendices………………………………………………………………………………………… 455





























v

Acknowledgements

First of all I would like to thank especially my mother and my father. Both of
them supported strongly this long term project, which at many times through the years
was not the easiest task to bear. Their love and understanding, always soothed the
difficult moments, and always boosted up my broken motives. This thesis, first and
above all, is dedicated to you.

Then I sincerely want to express my deep appreciation for the director of my
thesis, Jose Salvador Peris Alvarez. Salvador, by accepting me to the University of
Salamanca, you let me in into a world of knowledge, science and experiences that I
could not have imagined before. So firstly, I thank you for receiving me as a graduate
student from Greece, and transforming me into a Doctor. And secondly, because
through this process, I came to Spain, lived there for years, I got to know its everyday
life, I met its people, I made loving, lasting friendships, and that way this country,
became my second home. A priceless, unique, lifetime’s experience. I will always
owe you, Salva.

There are two more professors in Greece, to whom I owe a great deal. Christos
Vlachos and Dimitrios Bakaloudis. You were my University teachers and my first
mentors. From you I firstly heard the meanings Wildlife Biology, Sustainable
Management, Raptor Ecology. With you I firstly visited the biodiversity hot spots of
Greece, to work as a volunteer; with you I firstly walked the path of investigation, and
exhausting data field collection; with you I firstly experienced the unforgettable
feeling, of working as a team in the field, and although tiredness was always adding
up as the sun went down, I never wanted the day to end. Christos, exactly 10 years
ago in the village of Stefanovikeio, you showed me the first pair of Barn owls that I
had ever observed in the field. I’ll never forget the feeling. I guess, by completing this
thesis, I demonstrate among other things the impact that these moments had on me.
Dimitris, I recall you with your father in the river of Evros, in the frontier with
Turkey, recognizing raptors without binoculars, just by the way they were flying over
us, and arguing, who was right, and who was wrong. All I can wish now is that
projects will keep coming up!

During the realization of this thesis, two more people played a definitive and
very essential role for its completion. Ignacio Torre Corominas and Jaime Madrigal
Gonzalez. Ignacio, with your deep knowledge on small mammal behavior and
ecology, and your long term recognized investigation in this field, you offered me
great insight and new perspectives for my thesis, while at the same time your critic
and statistical advices unblocked the path for me at very difficult times. Many thanks,
and deep appreciation. Jaime, I will never forget these four days in Cuellar. You
introduced me to the world of multivariate analysis of ecological data, a scientific
field that you dominate, in which you are an expert. And your wonderful mother,
calling us to lunch and dinner, for quick breaks away from our computer screens. You
are an excellent investigator, and you offered me great help. But most of all, you are a
dear and supporting friend. I thank you for the first. And I feel blessed and lucky for
the latter. May your family, you and Tere, be always well, wherever your paths may
lead you.

vi

For the collection of data concerning the study area (the lowlands of
Thessaly), a great deal of public services, offices, organizations, NGOs, ministries and
investigation centers were visited. During these multiple visits, which quite often
were proved to be very disappointing and frustrating, there were a few people who
offered a great deal of help and I feel the need to thank them. In the Statistical Service
of Larissa, the director in charge, Ms Rizou, was very kind and helped a lot with
providing data. In the National Agricultural Research Foundation of Larisa, the
investigators Leonidas Toulios, Margaritis Toulios and Tilemaxos Lelentzis, provided
excellent help, maps and data, but above all their knowledge in edafological aspects
of the region. Miguel Lizana, who was one of my professors during the first two years
of my Phd studies in the University of Salamanca. He supported strongly my goal,
and in certain difficult times his help was more than valuable. And finally Vasileios
Goutner, who is a Full Professor in Aristotle’s University of Thessaloniki in the
Biology Department. He was a pioneer in publishing important Barn owl diet studies
during the recent years in Greece, and he repeatedly accepted me in his office, to
analyze with me various aspects of my thesis and answer my questions.

I also am deeply indebted to the Spanish Ministry of Foreign Affairs and the
Department of International Cooperation, for granting me a scholarship in the
beginnings of my Phd studies in Spain, and to Chloros Foundation and the National
Metsoveio University, for granting me an additional scholarship to complete my
research and Phd thesis.

Last but not least, I also wish to thank two more groups of people. At first, my
dear friends in Greece. Real, considering, unique friends, who each one with his own
special way, walked along with me this path that I chose, and was always there to
share a laugh, a thought, a glass of wine, a tear, a moment. First of all Afroditi, who
apart from being my sister, was also a real friend, always, in all circumstances. Sofia,
Giannis and Themis, who always gave me the last hug before each trip to Spain, and
always expected me back in Athens with a bottle of wine in the table. Giannis, Dora,
Vasilis, Giorgos, Aggeliki and Fotis who live in Thessaloniki, the city where
everything started, and will always be my favorite, as long as you are part of it. And
finally in Larisa, my hometown, Achilleas and Aliki with her family. I love you all.

Secondly, I sincerely want to thank, all these people that I met during my three
year field investigation, in the vast region of Thessaly. They changed the way I see
things, life, they broadened my perspective, and they were always there, to bring a
glass of water, to a sweating, strange guy with a helmet, who was visiting forgotten
villages, searching all these abandoned houses, ready to collapse. Without trying, they
showed me the other face of life; they showed me my land, my home, and I learned to
appreciate it even more, through their eyes.

This Phd thesis is the fruit of a long race. Finally, I just wish, that it can be of
some use, as a small help for others to step on, and reach beyond, even further, in this
specific field of science. In Thessaly, in Greece, or in any other part of the world…





vii

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domingo, 09 de junio de 2013 - 1:37