Ulfc103114 TM Margarida Goncalves PDF

UNIVERSIDADE DE LISBOA
FACULDADE DE CIÊNCIAS
DEPARTAMENTO DE BIOLOGIA ANIMAL
Phylogenetic diversity of bird assemblages along different land-use

gradients: regional and continental analysis
Margarida Ladeira Felício Gonçalves
Dissertação
Mestrado em Biologia da Conservação
2013
UNIVERSIDADE DE LISBOA
FACULDADE DE CIÊNCIAS
DEPARTAMENTO DE BIOLOGIA ANIMAL
Phylogenetic diversity of bird assemblages along different land-use

gradients: regional and continental analysis
Margarida Ladeira Felício Gonçalves
Dissertação orientada por Professor Dr. Roland Brandl, Departamento de

Ecologia,Phillips-Universität Marburg e Professor Dr. Carlos Fernandes, Departamento
de Biologia Animal, FCUL
Mestrado em Biologia da Conservação
2013
Dedicatória
À minha mãe.
Embora ausente, o teu amor é a minha força, iluminando o meu
caminho.
Thankful note/Agradecimentos
Primarily I would like to thank PhD Prof. Roland Brandl from Philipps-
Universität Marburg for accepting me as his Master student, accompanying me
and teaching me throughout my ERASMUS placement. For his patience, total
availability, tolerance and dedication put upon my work.
To his workgroup, a special thanks to Eugene Egorov for all the patience,
data and information provided and to Maike Franzen for all the co-work,
company and friendship.
Um agradecimento ao Prof. Dr. Carlos Fernandes, da FCUL, por me ter
aceitado como sua orientanda, pela disponibilidade e dedicação, sobretudo nos
últimos meses de trabalho.
À sua equipa, queria agradecer pessoalmente ao André Silva pela
disponibilidade e prontidão.
Ao meu Pai, pela força, esperança e amor. Por acreditar sempre em mim
e no meu trabalho.
Aos meus irmãos, sobrinhos, cunhados e tios-avós pela força, apoio e
preocupação.
A todos os meus amigos que sempre me apoiaram.

Contents
Summary ............................................................................................................ 1
Resumo .............................................................................................................. 3
1. Introduction .................................................................................................. 7
1.1 Phylogenetic diversity and conservation ............................................... 7
1.2 Processes structuring assemblages ...................................................... 8
1.3 Effect of anthropogenic land-use on phylogenetic diversity................... 9
1.3.1 Agriculture .................................................................................... 10
1.3.2 Urbanization.................................................................................. 11
1.4 Aims .................................................................................................... 12
2. Methods ..................................................................................................... 14
2.1 Data collection ..................................................................................... 14
2.2 Phylogeny ........................................................................................... 15
2.3 Spatial distribution of species .............................................................. 16
2.4 Phylogenetic structure of assemblages ............................................... 16
2.5 Effects of anthropogenic land-use on phylogenetic diversity ............... 17
2.5.1 Definition of land-use, bioclimatic and trait variables .................... 17
2.5.2 Generalized Additive Models ........................................................ 22
3. Results ....................................................................................................... 25
3.1 Spatial distribution of species .............................................................. 25
3.2 Phylogenetic structure of assemblages ............................................... 26
3.3 Effects of anthropogenic land-use on phylogenetic diversity ............... 26
4. Discussion ................................................................................................. 41
4.1 Spatial distribution and phylogenetic structure of assemblages .......... 41
4.2 Anthropogenic land-use impact ........................................................... 42
4.3 Implications for conservation ............................................................... 48
5. References ................................................................................................ 50
6. Appendix .................................................................................................... 59
Summary
Land-use intensification leads to the transformation of natural habitats
into antrhopogenic ones, such as farmland and meadows or, in its widest
sense, into urban areas. These habitats act as environmental filters, selecting
only those species whose traits enables them to survive, and although a
potential species richness increase, such species belong only to a set of a few
lineages. Following an evolutionary conservative approach, it is thus expected a
decrease of phylogenetic diversity of assemblages with increasing amount of
transformed habitats.
The present study evaluates the phylogenetic diversity (mean
phylogenetic and mean nearest neighbour distance) of native breeding bird
assemblages, considering the impact of land-use at regional and continental
scale, using Generalized Additive Models (GAMs); species richness was used
for comparative reasons only. Regionally, across Bavaria, it was used
presence/absence maps within 34 km2 grids and different land-use types; in a
European-wide study it was analysed which factors contribute to this
assemblage composition, in general and which of them differ between natural
and antrhopogenic sites.
Overall it was found that the occurrence of bird species is driven by
habitat filtering and not competition, regardless the scale of analysis. Within
regional analysis although species richness increased with increasing
percentage of anthropogenic land-use, both measures of phylogenetic diversity
showed an overall decrease. For European-wide analysis, overall diversity was
positivivelly related to area and climate. Natural and anthropogenic sites show
1
no significant differences, except only for species richness regarding body mass
(g) and percentage of insectivores.
In a world where urbanization is increasing at an exponential rate it is
important to analyse and recognize what processes and which factors
contribute the most for shaping animal assemblages. These results confirm the
elevated impact that anthropogenic habitats, especially urban areas, have upon
shaping bird assemblages, acting as strong environmental filters, with a
powerful evolutionary force.
Key-words: phylogenetic structure, habitat filtering, urbanization, Generalized
Additive Models
2
Resumo
A diversidade filogenética foi primariamente aplicada ao estudo da
conservação, sendo que as espécies prioritárias eram seleccionadas com base
na sua taxonomia. Hoje-em-dia, a importância da aplicação desta medida de
biodiversidade extende-se também ao estudo das comunidades, promovendo o
conhecimento dos processos evolutivos que levam à composição da sua
estrutura filogenética, bem como da interacção existente entre as espécies
constituintes. Dois processos são vistos como centrais nesta temática: (1)
exclusão competitiva entre espécies que limita a sua coexistência a longo
prazo; e (2) filtragem ambiental de espécies que persistem numa comunidade
devido à sua tolerância a factores abióticos. O primeiro processo leva a uma
sobre-dispersão filogenética, que tende a limitar a coexistência de espécies
relacionadas entre si, conduzindo a uma diversidade filogenética superior à
esperada sob condições aleatórias de estrutura comunitária. Pelo contrário, o
segundo processo leva a um agrupamento filogenético, no qual espécies
relacionadas tendem a coexistir e a diversidade filogenética revela-se menor
que o esperado.
A intensificação do uso-da-terra tem, nas últimas décadas, levado à
transformação do ambiente em habitats antropogénicos, nomeadamente em
terrenos agrícolas e de pastoreio, ou num sentido mais extremo, em áreas
urbanas. É do conhecimento geral que tais habitats altamente modificados
actuam como filtros ambientais, selecionando apenas algumas espécies com
determinadas características que lhes permitem sobreviver e que pertencem
assim, a apenas deteterminadas linhagens. Seguindo um contexto evolutivo
3
conservacionista espera-se que, em áreas antropogénicas as espécies tendam
a coexistir, levando à diminuição da diversidade filogenética, mesmo quando
um aumento da riqueza específica se poderá também verificar.
Como tal, o presente estudo avalia a diversidade filogenética de
comunidades de aves nativas nidificantes, considerando o impacte do uso-da-
terra, tanto a nível regional como a nível continental. A diversidade filogenética
foi calculada usando duas medidas diferentes: distância filogenética média,
mais sensível a padrões que se verificam a nível geral, em toda a árvore; e a
distância média ao vizinho mais próximo, mais sensível a padrões que ocorrem
nas extremidades da árvore filogenética. Para a análise foram usados modelos
estatísticos, nomeados Modelos Aditivos Generalizados (GAMs), nos quais a
função de ligação dos Modelos Lineares é substituída por uma função não
paramétrica, estimada através de curvas de alisamento que permitem
descrever a forma da função e revelar possíveis não linearidades nas relações
estudadas. A riqueza específica foi também incorporada, apenas para termos
comparativos.
Regionalmente, o estudo foi efectuado no estado alemão da Baviera,
usando dados de presença/ausência de espécies, recolhidos entre 1996 e
1999 em mapas com grelhas de cerca de 34 km2 e testando a influência da
percentagem de diferentes tipos de uso-da-terra. A nível continental, foram
usados estudos de presença/ausência documentados em vinte-e-dois países
europeus entre 1973 e 2008, testando a influência da área de estudo e
variação temporal na comunidade de aves europeias, bem como os diferentes
factores que contribuem para a composição desta comunidade, em geral e os
4
que induzem a possíveis disparidades que se observam na composição entre a
comunidade existente em ambientes naturais e a comunidade antropogénica.
Os resultados provenientes da distribuição espacial das espécies
demostram que a ocorrência das espécies de aves analisadas é determinada
por filtragem ambiental e não exclusão competitiva, independentemente da
escala analisada. Isto leva a um processo de agrupamento filogenético e a uma
menor diversidade filogenética do que seria de esperar sob condições neutrais.
Na Baviera, como esperado, apesar de registado um aumento da riqueza
específica com o aumento da percentagem de todos os tipos de uso-de-terra
definidos, foi igualmente verificado um decréscimo da diversidade filogenética,
para ambas as medidas calculadas. A nível europeu a riqueza específica e
ambas as medidas de diversidade filogenética foram positivamente
relacionadas com o aumento da área de estudo e clima, nomeadamente o
aumento da temperatura. Contrariamente ao esperado, não foram detectadas
diferenças significativas a nível filogenético entre ambientes naturais e
antropogénicos; estas diferenças só ocorreram para a riqueza específica
relativamente a duas variáveis: massa corporal (g) e percentagem de espécies
insectívoras.
Estes resultados mostram, primariamente, a importância de incorporar
não só diversas medidas de diversidade, bem como diferentes escalas de
observação em estudos relativos à análise e gestão de comunidades. Mostram
também o impacte que diferentes tipos de uso-da-terra, nomeadamente zonas
urbanas, têm na organização e estruturação destas comunidades, actuando
como ambientes filtradores que seleccionam as espécies com características
que lhes permitem sobreviver e adaptar à presença constante do Homem. A
5
nível europeu, os resultados enfatizam a importância que áreas contínuas e de
grande dimensão têm na gestão das comunidades de aves, sobretudo em
zonas com condições climáticas favoráveis à reprodução e nidificação destas
espécies, como por exemplo, em países do sul da Europa. Contrariamente ao
esperado, não foram detectadas diferenças significativas entre comunidades
europeias naturais e antropogénicas, no espaço temporal analisado; apenas
foram detectadas diferenças em relação à riqueza específica, com maior
número de espécies de menores dimensões e aumento do número de espécies
com uma maior percentagem de aves insectívoras em zonas antropogénicas. A
longo prazo, este processo selectivo poderá levar a um fenómeno de
homogeneização biótica, intimamente ligado à perda de biodiversidade a nível
global.
Este estudo promove, assim, a aplicação de acções de gestão que
mantenham as espécies existentes e que aumentem a diversidade filogenética
das comunidades analisadas, podendo os resultados ser abrangidos a outras
regiões com características semelhantes, assim como a outros taxa existentes.
Num futuro onde a urbanização crescerá a níveis exponenciais, é importante
perceber as condições que estes habitats proporcionam e o potencial oferecido
em termos de biodiversidade.
Palavras-chave: estrutura filogenética, filtragem ambiental, urbanização,
Modelos Aditivos Generalizados
6
1. Introduction
1.1 Phylogenetic diversity and conservation
Understanding the mechanisms and processes that shape the
distribution of phylogenetic diversity along environmental gradients can become
crucial to the study of the potential effects of global change, identification of
vulnerable ecosystems or species and the proposal of meaningful conservation
measures to mitigate the current diversity crisis (Winter et al. 2013). Successful
conservation strategies should thus retain as large an amount of phylogenetic
diversity as available resources permit (Faith et al. 2004) and this parameter
can be targeted directly into conservation planning (Rodrigues & Gaston 2002;
Rolland et al. 2012).
Faith (1992) was among the first researcher that quantified phylogenetic
diversity (PD) as the cumulative length of the branches connecting the root of
the phylogenetic tree, representing the common ancestral lineage, to the
evolving tips, represented by species. Phylogenetic branch lengths are a
continuous metric of relatedness and this measure was primarily applied to
conservation, where the priority taxa to be protected reflected a certain value of
biodiversity (Minh et al. 2006). With the increasing availability of phylogenetic
information it is now possible to develop more sophisticated methods to infer
processes that structure the composition of species assemblages at different
scales (Pavoine et al. 2005; Webb et al. 2002).
Conservation research has been primarily focused on a global-scale of
phylogenetic diversity loss; however the loss of PD at smaller spatial scales is
7
of equal concern, since losing diversity at any scale can lead to a reduced
potential for assemblages to respond to changing environmental conditions,
through a reduction of genetic diversity (Morlon et al. 2011). Also, assemblages
share a greater fraction of PD than species, suggesting that single isolated
areas can be more efficient at preserving phylogenetic diversity than species
richness (Swenson 2009).
1.2 Processes structuring assemblages
Species’ features are known to influence their interaction with other
species and with the environment. If, on the one hand, phylogenetically closely
related species are ecologically similar and therefore tend to co-occur, it is also
expected that related species compete with each other, which constrains their
coexistence (Losos 2008). In this way, depending on scale, bird assemblages
are structured by a series of processes that can range from neutral or niche-
based processes at small scales to speciation and global extinction at larger
scales (Cavender-Bares et al. 2009). Such processes are expected to leave a
distinct signal in the phylogenetic structure of assemblages (Cadotte et al.
2010) since ecologically similar species also tend to be phylogenetically related
and therefore ecological factors of assembly composition are reflected in
phylogenetic patterns (Helmus et al. 2007).
Niche-based processes may be divided into competitive interactions and
environmental filtering. Assuming that phylogenetically related species occupy
similar niches, competitive interactions should lead to assemblages in which
species are phylogenetically less related (phylogenetic overdispersion) whereas
8
environmental filtering should lead to assemblages in which species are
phylogenetically more related (phylogenetic clustering) when compared to
assemblages solely structured by neutral processes (Burns & Strauss 2011;
Emerson & Gillespie 2008; Kluge & Kessler 2011).
About 60 years ago researchers started to use the species to genus ratio
as a simple measure of phylogenetic structure to search for competitive
interactions, a niche-based process (Elton 1946). These competitive
interactions should be particularly important between closely related species
and therefore assemblages structured by competition should lead to a lower
species to genus ration than expected by chance (Mayfield & Levine 2010).
The importance of the various processes that structure a community
depends on spatial (and temporal) scale (Devictor et al. 2010). By previous
studies (Vamosi et al. 2009) it is known that competitive interactions may
become important on small scales whereas habitat filtering may be important on
larger scales. Nevertheless, in a recent study (Gotelli et al. 2010) it has been
shown for birds across Denmark, that competitive interactions leave a signal in
the co-occurrence of species analysed on a grain size up to four orders of
magnitude larger than the territory size, slightly contradicting previous
statements.
1.3 Effect of anthropogenic land-use on phylogenetic diversity
Beside biotic processes, the most important determinant of specie’s
distribution is the scattering of habitat across a landscape (Franklin 2009;
Peres-Neto et al. 2012). Furthermore, the characteristics, distribution and area
9
of the various habitats are influenced by human activities, thereby changing the
habitat that is available for the species across that landscape (Butler et al.
2010).
Anthropogenic land-use, defined as special human created habitats,
such as farmland, meadows and urban areas, covers growing proportions of the
global landscape (Filippi-Codaccioni et al. 2010). Particularly, agriculture and
urbanization lead to fragmentation of natural areas, offering a set of special
conditions for birds and other wildlife species by modifying processes like
predation, interspecific competition and diseases, which structure bird
assemblages. Only a subset of species may be able to cope with such special
conditions (Alberti 2005). Consequently, one of the most recognized
disturbances that may lead to niche-based processes in assemblages is human
changed land-cover and land-use (Fuller & Gaston 2009). However, it is also
known that anthropogenic habitats can accommodate an astonishingly rich flora
and fauna, which can achieve large population sizes (Chiari et al. 2010; Hole et
al. 2005; Rodewald & Shustack 2008).
1.3.1 Agriculture
Agriculture is considered the most important type of land use in Europe;
however, nowadays it is also acknowledge that agricultural intensification is the
major cause of decline in the abundance and diversity of bird species in this
continent since the 1970’s (Donald et al. 2001). This intensification has led to
the fragmentation and simplification of habitat, with increasing monocultures
and fewer non-cultivated land, due to increased use of agrochemicals and
10
machinery, which reduces food and nest availability for birds (Batáry et al. 2007;
Belfrage et al. 2005).
Nevertheless, a range of birds are still known to depend on agricultural
land, especially in winter (Atkinson et al. 2005) and more than 50% of the
existing habitats in Europe occur in farmland (Söderström et al. 2001),
supporting more bird species of conservation concern than any other habitat
type (Wilson et al. 2005). This is possible because throughout European
countries one can find different farming systems and management practices
[e.g. organic vs. conventional farming (Darnhofer et al. 2010; Fuller et al. 2005);
set-aside land (Buskirk & Willi 2004); and agri-environment schemes (Donald &
Evans 2006)], which result in a vast spectrum of specie’s responses (Reidsma
et al. 2006).
1.3.2 Urbanization
Urbanization is a complex process of land conversion and it is now
considered the fastest growing land-cover type, showing an exponential growth
in Europe since the end of the 19th century (Antrop 2004). This process affects
bird species through profound and permanent changes in the habitat (Bowman
& Marzluff 2001).
However, much like agriculture, cities may also benefit birds through
higher resource abundance, lower predator pressure (Jokimäki & Huhta 2000;
Shochat et al. 2010), ameliorated climatic conditions, increased water
availability and increased habitat availability that results from edge effects and
ornamental vegetation (Marzluff 2005). Therefore, considered at the landscape
11
scale, cities may have a positive effect on species richness (Kühn et al. 2004;
Mehr et al. 2011), especially due to an increasing adaptation from exotic
species (Loss et al. 2009).
1.4 Aims
In a previous study, Pfeifer et al. (2009) evaluated bird species
assemblages across Bavaria and discovered that species richness increased
with increasing proportion of anthropogenic habitats, such as cities or industrial
areas, when species richness was evaluated on a grain of approximately 34
km2. Similarly, a study on bats for the same area found also an increase of
species richness with increasing percentage of anthropogenic habitats; however
a decrease of phylogenetic relatedness between species (Riedinger et al.
2013). This leads to the idea that such modified habitats may have contrasting
effects on species richness and phylogenetic diversity.
Following this, the main goal of the present study is to understand what
phylogenetic signals anthropogenic habitats leave on native breeding bird
assemblages, when focusing on different scales. At the regional scale across
Bavaria on a grain of 34 km2 and by using broad habitat types, labelled land-
cover types; at the continental scale across Europe, along a spatial and
temporal gradient, using a grain from 1.04 and 108 333 km 2 during thirty-five
years, and also testing which factors contribute to significant differences
between natural and anthropogenic assemblages.
More specifically it aims to 1) test the hypothesis that competitive
interactions influence the distribution of native breeding bird species across
12
Bavaria, in particular and Europe, in general; 2) evaluate the phylogenetic
structure of both assemblages, determining if they are more (phylogenetic
clustering) or less related (phylogenetic overdispersion) than expected when
compared with an assemblage structured by neutral processes; 3) test the
hypothesis of an expected decrease of phylogenetic diversity with increasing
proportion of anthropogenic habitats at regional scale; 4) evaluate wich factors
contribute to the phylogenetic structure of European’s bird assemblage in
general; and 5) test the hypothesis that natural and antrhopogenic areas within
Europe present significant differences regarding some or all of these factors.
13
2. Methods
2.1 Data collection
Data collection was performed differently for both parts of the study,
since they differ in the scale of analysis. Both regional and continental studies
were thought considering the information available for bird community ranges
and data resolution concerning land-use types, climate and ecological variables.
Regional analysis was performed in Bavaria, a federal state in the south
east of Germany, covering an area of 70 547.82 km², half of which is classified
as agricultural land. Annual mean temperature varies from 14.4 ˚C and 2.1 ˚C
and mean annual precipitation between 850 mm and 1000 mm (Bezzel et al.
2005). Bird’s distribution originated from the same mapping project used by
Pfeifer et al. (2009), using grid maps with information recorded between 1996
and 1999 collected by Bavaria’s Environmental State Secretary, Ornithology
Station Garmisch-Partenkirchen (Bezzel et al. 2005). Each grid covered an
average area of 33.9 km2 (minimum 32.9 km2; maximum 35.1 km2), and only
grids fully within the Bavarian borders were used (1927 grid cells out of a total
of 2285 – Appendix A). The presence/absence of native breeding bird species
was recorded across these grids and only those who represented > 5% of total
observations were used for the analysis, with the exception of those whose
genera wasn’t represented yet, so that all genera recorded were present in the
data (in total 157 species - Appendix B).
Continental analysis was achieved with a total of 127 studies reporting
the presence/absence of bird species from 22 European countries collected
14
between 1973 and 2008 (Appendix C). Species were only selected if
considered native and breeding within the country where they were present.
Studies were categorized according to the level of human impact/urbanization
of the site where they were performed: pristine sites to small fields, including
several types of habitats (e.g. mountains, river sides and forests) were
classified as Natural (80 studies); agricultural areas and villages with more than
1 000 inhabitants to capital cities were classified as Anthropogenic areas (47
studies). Species selection was the same as for Bavaria: only species with more
than 5% of total observations were used for the analysis, with the exception of
those whose genera wasn’t represented yet, so that all genera recorded were
present in the data (in total 297 species - Appendix D).
2.2 Phylogeny
The basis for the evaluation of phylogenetic relatedness within
assemblages was a phylogeny retrieved from a global phylogeny of birds (Jetz
et al. 2012b). For both assemblages studied it was obtained a phylogeny
corresponding to the species sampled in each matrix data, after the following
process. First, for the species recorded in each assembalge a set with 100
phylogenies was retrieved from www.birstree.org using the Ericson All Species
Source Tree [10 000 trees covering 9 993 species each (Jetz et al. 2012a)]. For
each tree a distance matrix was calculated between species using the function
distTips in the add-on package adephylo in R (Jombart 2013) and after, the
mean distance across all 100 matrices. These matrices constitute an estimate
of the phylogenetic distance between species and the distance was about twice
15
to the common ancestor. The final tree for each phylogeny (Appendix E & F)
constituted the hierarchical cluster analysis of the previous calculated mean,
using the un-weighted average linkage algorithm.
2.3 Spatial distribution of species
In order to test the hypothesis that competitive interactions influence the
distribution of species for both assemblages, it was used the function
comm.phylo.cor in the add-on package picante in R (Kembel 2010) with 999
randomizations to evaluate significance. This function characterizes the
distributional overlap across a community, checking for a correlation between
co-occurrence patterns of species and phylogenetic distance between them.
This index is based upon Schoener’s index, originally used to measure
overlap in habitat use or diet (Schoener 1970). An index of 0 indicates that two
species never occur together in the same grid and an index of 1 indicates that
the two species always occur together. Other available indexes were used
(Jaccard, Checkerboard and DOij indexes), since they only differ by the way co-
occurrence is estimated and therefore provide almost the same results (Hardy
2008).
2.4 Phylogenetic structure of assemblages
After this, and also using the add-on package picante in R, it was
evaluated the phylogenetic structure of the assemblages, using the matrix of
average phylogenetic distance and by calculating the standardized effect size of
16
the mean phylogenetic distance (SESMPD) and of the mean nearest neighbour
distance (SESMNTD), whose functions are as follows:
( )
where index.obs is, respectively, the observed mean phylogenetic distance or
mean nearest neighbour distance in communities; index.rand.mean is the mean
distance calculated from a null model and index.rand.st is the standard
deviation of the index across null communities. Values < 0 indicate phylogenetic
under-dispersion (clustering) whereas values > 0 indicate phylogenetic over-
dispersion. The null model was obtained by reshuffling species names across
the tips of the phylogeny, using 999 runs. This model keeps the occurrence, as
well as recorded number of species within grids unchanged.
2.5 Effects of anthropogenic land-use on phylogenetic diversity
2.5.1 Definition of land-use, bioclimatic and trait variables
To test for the hypothesis of a decrease phylogenetic diversity with
increasing proportion of anthropogenic land-use types at regional scale and
evaluate the factors that contribute to European assemblage as well as for
differences between natural and antrhopogenic sites, it was analysed the
response of the effect size of the mean phylogenetic distance as well as the
effect size of the mean nearest neighbour distance to different land-use
variables and both area and time variation. It was also analysed the response of
17
space (Latitude and Longitude), bioclimatic and trait variables as control. This
process was achieved using Generalized Additive Models – GAMs.
For Bavaria, it was used the same land-use (habitat) and climate data as
both Pfeifer et al. (2009) and Riedinger et al. (2013). Data included one set that
characterized the climate, extracted from the 19 bioclimatic variables available
in WorldClim (http://www.worldclim.org) using a spatial resolution of 30
seconds. The second set of data consisted of percentages of area covered by
broad habitat types in each grid and its location (Latitude and Longitude). This
land-cover was derived from the land-cover classes after the European wide
mapping project CORINE (Coordinated Information of the European
Environment), which uses LANDSAT-7 satellite images (scale 1:100 000)
collected in 2000 (http://www.corine.dfd.dlr.de). Several classes were grouped
into seven broad land-use types (Table 1), which were used in subsequent
analysis. Both data sets were read into ArcGis 10.2.
For continental analysis each study was characterized by both a spatial
compoent, representing the dimension of the study-area in km2, with a minimum
area of 1.04 km2 and a maximum area of 108 333 km 2 and a temporal
component, represented by the average year of each study, along thirty-five
years. Land-use was performed by quantatively selecting the study-site as
natural or anthropogenic, as mentioned earlier and climate was analysed using
only 4 of the 19 bioclimatic variables from WordClim, since only these were
thought to best define the seasonality differences that occur intra-continent
(bio5, bio6, bio13 and bio14). Climate data was also read into ArcGis 10.2.
18
Table 1. Land-use types formed from the CORINE land-cover classes. For each class it was
given the code used by the CORINE web site.
Land-use types Code CORINE land-cover classes

211 Non-irrigated arable land
Farmland 221 Vineyards
222 Fruit trees and berry plantations
231 Pastures
Meadow 243 Agricultural land with areas of natural vegetation
321 Natural grassland
322 Moors and heathland
411 Inland marshes
412 Peat bogs
Wetland
331 Beaches, dunes, sand
511 Water courses
512 Water bodies
Deciduous forest 311 Broadleaf forest
Coniferous forest 312 Coniferous forest
Mix forest 313 Mix forest
121 Industrial, commercial and public units
124 Airport
131 Mineral extraction sites
132 Dump sites
111 Continuous urban fabric
Anthropogenic habitats 112 Discontinuous urban fabric
123 Port areas
133 Construction sites
141 Green urban areas
142 Sport and leisure facilities
122 Road and rail networks and associated land
For both assemblages it was also added a trait set characterizing
biological variables for each species analysed, including body mass (g), trophic
niche, clutch size (Cramp et al. 1994; Dunning 1993) and population size
19
(retrieved from http://www.iucnredlist.org and http://birdlife.org) since all of them
contribute for specie’s sensitivity to human impact and consequent priority for
management planning. All of these variables, although specie’s specific were
converted into assembalge based, in order to minimize within site/study
variation due to specie’s trait differences. This was accomplished by calculating
the logarithmic mean of body mass (g), clutch size (per year) and population
size (pairs x 10000), averaged across all species that were present on each site
or study and average percentage of species with each diet category (e.g.
carnivores) (Table 2 & 3).
Table 2. Trait variables analyse for Bavarian assemblage. SD stands for standard deviation;
also included the 25% and 97.5% range of each trait sample.
Bavaria
Mean ± SD 25% 97.5%

Body mass (g) 48.36 ± 9.446 42.439 67.046
% Carnivores 9.890 ± 2.899 7.828 15.058
% Insectivores 44.46 ± 3.629 42.169 51.341
% Omnivores 27.17 ± 3.223 25 34.311
% Piscivores 1.583 ± 1.380 0 4.440
% Granivores 16.890 ± 2.605 15.068 22.346
% Scavengers 0.295 ± 0.605 0 1.754
Clutch size (year) 5.980 ± 0.228 5.835 6.456
Population size (pairs x 1000) 4.083 ± 1.720 3.421 6.365
20
Table 3. Trait variables analyse for natural and anthropogenic sites within European-wide
assemblage. SD stands for standard deviation; also included the 25% and 97.5% range of
each trait sample.
European natural sites European anthropogenic sites
Mean ± SD 25% 97.5% Mean ± SD 25% 97.5%

Body mass (g) 73.99 ± 42.078 52.374 163.589 50.28 ± 13.070 41.761 75.041
% Carnivores 10.5 ± 4.060 8.649 18.193 8.575 ± 2.881 5.742 12.617
% Insectivores 49.17 ± 4.848 45.213 60.031 47.13 ± 4.933 43.584 52.549
% Omnivores 23.98 ± 4.132 20.572 31.119 26.25 ± 4.361 23.181 35.635
% Piscivores 3.143 ± 2.890 1.127 12.071 1.579 ± 1.364 0 4.85
% Granivores 13.880 ± 3.704 11.746 21.966 15.840 ± 3.967 12.596 22.228
% Scavengers 0.710 ± 0.760 0 2.592 0.189 ± 0.350 0 0.959
Clutch size (year) 5.633 ± 0.415 5.384 6.333 5.830 ± 0.252 5.652 6.256
Population size (pairs
37.850 ± 26.929 19.702 100.720 3.513 ± 0.906 2.848 5.213
x 1000)
Since the bioclimatic set contained a large amount of data and variables
were highly correlated, this set was summarized with principal component
analysis based on the correlation matrix for each assemblage and then the first
two principal components where used for further analysis (Appendix G). Both
components summarized together an average of 70% of the total variance. In a
general way, PC1 is a measure of annual temperature and precipitation
whereas PC2 measures the seasonal variability of the climate.
Before anyfurther analysis it is important to test the existence of
collinearity between response variables in order to prevent weak results and
facilitate variables’ relation prediction (Dormann et al. 2013). Collinearity was
tested with simple pairwise correlation coefficient (r) for each variable within
each assemblage and the threshold was set at r > 0.7.
21
For both Bavarian and European assemblages it was found a high
correlation between Latitude and PC1 (Bavarian r = 0.71; Anthropogenic sites r
= 0.85; P < 0.001) and clutch size and population size (Bavarian r = 0.77;
Anthropogenic sites r = 0.99; P < 0.001), so both Latitude and clutch size were
removed from both data. Only for Europe, the percentage of Piscivores and
body mass were highly related (r = 0.75; P < 0.001), so the first variable was
removed from this data.
2.5.2 Generalized Additive Models
Generalized Additive Models – GAMs – were first introduced by Hastie &
Tibshirani (1986) and are extensions of Generalized Linear Models (GLMs)
where the linear predictor is given by a specified sum of smooth functions of
covariates. It can be simply defined as follows:
( ) ( ) ( )
where the response variable yi can be defined by smooth functions k1 and k2 of
covariates x1 and x2. It predicts some known smooth monotonic function of the
expected value of the response variable, which may follow any exponential
family distribution, permitting the use of a quasi-likelihood approach (Wood
2006).
The effect sizes of the mean phylogenetic distance and of the mean
nearest neighbour distance were used as response variables for statistical
analysis using function gam from package mgcv in R (Wood 2013). For
22
comparative reasons it is also used the value of species richness. This package
solves the smoothing parameter estimation problem using the Generalized
Cross Validation (GSV) criterion, based upon the number of data, deviance and
effective degrees of freedom of the implemented model. The lower the GSV
score the better the model, with higher percentage of deviance explained.
For Bavaria, land-use data, both principal components of the climate data
and trait data were used as independent variables; for European-wide
assemblage it was the same, except for the land-use data analysis, which was
characterized by a factor that indicated if a study belonged to the natural or
antrhopogenic data set and the addition of both spatial and temporal
components within each site. This method of Generalized Additive Models
allows fitting curvilinear relationships and for this analysis it was used thin plate
splines as smoothers, starting with the default settings (k = 10) for Bavaria and
decreasing it to 5 for European-wide assemblage to obtain more robust models,
since the number of replicates was lower. Also, for European analysis the land-
use factor was added to the model in two different ways: individually, without a
smooth function describing it; and as a by-function for each independent
variable, allowing testing for significant differences between data-sets, within
Europe.
Furthermore, trait data can sometimes present a high correlation with
phylogenetic measures and space data always shows spatial autocorrelation,
which compromises additionally hypothesis testing. To remove the first
correlation, it was increased the default of the dimension of the basis used to
represent the smooth terms for trait variables and to control for spatial
autocorrelation it was also included the factor space in the gam models [see
23
also Dormann et al. (2007)]. For Bavaria the default of the dimension used to
represent the smooth term was increased to 40 and for European assemblage
to 10.
Also, the available significance tests for the smoothers are only
aproximate, and so it was followed a conservative approach, accepting
significance only if P < 0.01.
24
3. Results
3.1 Spatial distribution of species
The correlation between the phylogenetic distance averaged across all
100 distance matrices of the bird species with their co-occurrence was
significantly negative for both assemblages (Bavaria matrix correlation = - 0.29;
European matrix correlation = - 0.22; P ≈ 0.001). This means that phylogenetic
related species tend to co-occur more than expected, regardless the scale of
analysis.
Calculating this matrix correlation using distance matrices of each of the
100 trees retrieved for each assemblage analysed, also showed a negative
correlation for all matrices, with similar average numbers (Figure 1).
Figure 1. Histograms showing the distribution of matrix correlations for all 100 trees for both Bavarian
(on the right) and European (on the left) assemblages.
25
3.2 Phylogenetic structure of assemblages
The effect size of the mean phylogenetic distance and of the mean
nearest neighbour distance were moderately correlated for Bavaria and
European-wide assemblages (Bavarian r = 0.19; P < 0.001; European-wide r =
0.28; P < 0.01; both not corrected for spatial autocorrelation). The effect size of
the mean phylogenetic distance also showed a moderate to low correlation with
species richness for both assemblages (Bavarian r = 0.12; P < 0.001; European
r = 0.18; P < 0.05; both not corrected for spatial autocorrelation).
Averaged across all grids and studies, the effect sizes of the mean
phylogenetic distance as well as of the mean nearest neighbour distance were
negative for both assembalges and differed significantly from zero (Bavarian
SESMPD = - 4.37; t = -138.7; P < 0.01; Bavarian SESMNTD = -1.24; t = - 42; P <
0.001; European SESMPD = - 5.31; t = - 24.2; P < 0.001; European SESMNTD = -
1.88; t = - 19.6; P < 0.001), indicating community clustering in respect to
phylogeny at both tree-wide and closer-to-tips phylogenetic scale, regardless of
scale.
3.3 Effects of anthropogenic land-use on phylogenetic diversity
Considering Bavaria, GAMs results showed a relationship between land-
use variables and both measures of phylogenetic diversity, as well as species
richness (Table 4). The effect size of the mean phylogenetic distance was
related to meadows and anthropogenic habitats, presenting a decrease with the
increasing percentage of both (Figure 2).
26
Table 4. Results from Generalized Additive Models performed for Bavarian assemblage using species richness and both measures of phylogenetic
diversity as response variables. Est. Std. represents the estimated standard deviation; Std. Error the estimated error. Edf represents the estimated
degrees of freedom for each independent variable; Ref. df the estimated residual degrees of freedom for each variable. For each response variable is
also presented the R-square adjusted value - R-sq (adj) –, the GSV score and the explained deviation of each model. Significant values (P < 0.01) are
highlighted. For the land-use types, highlighted F values show an increase of the response variable with that land-use type variable.
Species richness Mean phylogenetic distance Mean nearest neighbour distance
Est. Std. Est. Std. Est. Std.

t-value P t-value P t-value P
Std. Error Std. Error Std. Error
Intercept 72.315 0.199 361.700 < 0.001 -4.368 0.012 -362.900 < 0.001 -1.241 0.017 -74.200 < 0.001
Edf Ref. df F P Edf Ref. df F P Edf Ref. df F P
PC1 climate 3.378 4.287 15.260 < 0.001 1.000 1.000 2.681 0.102 1.000 1.000 20.227 < 0.001
PC2 climate 5.453 6.610 9.014 < 0.001 3.492 4.442 4.404 < 0.001 1.000 1.000 0.266 < 0.01
Space 22.755 27.640 3.473 < 0.001 8.919 11.156 20.976 < 0.001 2.374 3.039 3.765 0.010
0.738
Farmland 1.473 1.810 21.270 < 0.001 4.044 5.013 1.566 0.166 1.000 1.000 0.112
27
Species richness Mean nearest neighbour distance Mean phylogenetic distance
Meadow 1.000 1.000 44.479 < 0.001 3.880 4.834 8.384 < 0.001 3.438 4.313 2.891 0.020
Wetland 4.351 5.283 8.388 < 0.001 1.000 1.000 0.740 0.390 1.960 2.469 7.823 < 0.001
Coniferous
3.021 3.817 6.204 < 0.001 3.183 4.017 5.058 < 0.001 4.449 5.500 1.008 0.412
forest
Deciduos forest 1.588 1.972 6.447 < 0.001 1.012 1.024 2.802 0.093 2.120 2.659 3.099 0.032
Mix forest 1.000 1.000 4.364 0.037 1.000 1.000 23.676 < 0.001 2.982 3.737 1.989 < 0.001
Anthropogenic
1.077 1.150 89.362 < 0.001 1.000 1.000 39.900 < 0.001 4.126 5.124 4.338 < 0.001
habitats
Body mass 4.708 5.919 22.361 < 0.001 5.109 6.377 480.892 < 0.001 5.305 6.609 1.258 0.268
Pop. size 14.380 17.330 5.587 < 0.001 12.843 15.555 4.342 < 0.001 2.558 3.099 34.201 < 0.001
Carnivores 9.303 11.605 5.642 < 0.001 35.842 37.525 3.839 < 0.001 12.807 15.841 1.928 0.015
Insectivores 4.679 5.995 16.566 < 0.001 12.366 15.202 4.755 < 0.001 1.000 1.000 6.166 0.013
Omnivores 14.982 18.459 6.633 < 0.001 5.587 7.122 7.071 < 0.001 7.566 9.527 3.198 < 0.001
Piscivores 20.512 24.551 7.954 < 0.001 8.176 10.008 4.380 < 0.001 1.000 1.000 63.091 < 0.001
28
Species richness Mean nearest neighbour distance Mean phylogenetic distance
Granivores 12.376 15.215 5.236 < 0.001 10.348 12.801 2.481 < 0.01 2.565 3.481 1.438 0.222
Scavengers 8.715 10.566 21.144 < 0.001 3.795 4.630 25.324 < 0.001 3.344 4.054 5.698 < 0.001
R-sq (adj.) 0.724 0.854 0.228
GSV score 82.872 0.298 0.557
Dev. Explained 74.40% 86.30% 25.30%
29
Figure 2. Scatterplots of the Generalized Additive Model for Bavarian assemblage relative to the
effect size of mean phylogenetic distance versus percentage of anthropogenic habitats (on top)
and percentage of meadows (on bottom). Red line represents the fit evaluated at the mean of the
independent variable and blue lines represent the fitted lines of ± two times the standard error of
the independent variable.
The effect size of the mean nearest neighbour distance was related to
wetlands and anthropogenic habitats. It showed a moderate decrease with the
increasing percentage of the first and with the second presented a hump-
30
shaped relationship, where nearest neighbour distance increased slightly until
the percentage of anthropogenic habitats reached about 15% and beyond this
point it decreased (Figure 3).
Figure 3. Scatterplots of the Generalized Additive Model for Bavarian assemblage relative
to the effect size of mean nearest neighbour distance versus percentage of wetlands (on
top) and percentage of anthropogenic habitats (on bottom). Red line represents the fit
evaluated at the mean of the independent variable and blue lines represent the fitted lines
of ± two times the standard error of the independent variable.
31
Species richness increased with the increased percentage of wetlands,
meadows (Figure 4), farmland and anthropogenic habitats (Figure 5).
Figure 4. Scatterplots of the Generalized Additive Model for Bavarian assemblage relative to
species richness versus percentage of wetlands (on top) and percentage of meadows (on bottom).
Red line represents the fit evaluated at the mean of the independent variable and blue lines
represent the fitted lines of ± two times the standard error of the independent variable.
32
Influence of climate variables and of total or single trait variables were
always significant for all three response variables (Table 4).
Figure 5. Scatterplots of the Generalized Additive Model for Bavarian assemblage relative to
species richness versus percentage of farmland (on top) and percentage of anthropogenic
habitats (on bottom). Red line represents the fit evaluated at the mean of the independent
variable and blue lines represent the fitted lines of ± two times the standard error of the
independent variable.
33
Considering European-wide assemblage, Generalized Additive Models
results presented significant relationship between the spatial component and
the effect size of the mean phylogenetic distance, as well as with species
richness (Table 5), with increased phylogenetic diversity (Figure 6) and number
of species (Figure 7) with area (km2).
Figure 6. Scatterplot of the Generalized Additive Model for European-wide assemblage relative to the
effect size of mean phylogenetic distance versus area (log in km2). Red line represents the fit evaluated
at the mean of the independent variable and blue lines represent the fitted lines of ± two times the
standard error of the independent variable.
Climate was significantly related to all three response variables, with
increasing species richness and overall phylogenetic diversity with increasing
mean annual temperature and low precipitation (PC1). Space had no influence
in any of the variables; total or single trait variables were always significant for
all three response variables.
34
Table 5. Results from Generalized Additive Models performed for European-wide assemblage using species richness and both measures of
phylogenetic diversity as response variables. Est. Std. represents the estimated standard deviation; Std. Error the estimated error. Edf represents
the estimated degrees of freedom for each independent variable; Ref. df the estimated residual degrees of freedom for each variable. For each
response variable is also presented the R-square adjusted value - R-sq (adj) –, the GSV score and the explained deviation of each model.
Significant values (P < 0.01) are highlighted. Regarding the independent variables, highlighted F values show an increase of the response variable
with that independent variable. It is also showen the results of the land-use factor interaction with each independent variable, allowing a between
natural and anthropogenic sites interpretation.
Est. Std. Est. Std. Est. Std.

t value |t| t value |t| t value |t|
Std. Error Std. Error Std. Error
Intercept 123.785 5.257 23.546 < 0.001 -5.434 0.268 -20.270 < 0.001 -1.554 0.307 -5.065 < 0.001
Land-use 13.371 9.630 1.389 0.170 -0.371 0.404 -0.920 0.360 0.207 1.743 0.119 0.906
PC1 climate 3.626 3.879 9.450 < 0.001 3.307 3.705 4.032 < 0.01 1.000 1.000 13.468 < 0.001
PC2 climate 1.000 1.000 3.444 0.068 1.000 1.000 0.130 0.720 1.000 1.000 0.464 0.498
35
Space 2.566 3.181 2.305 0.081 2.682 3.340 1.019 0.390 1.893 2.360 1.534 0.214
Area 3.791 3.954 27.377 < 0.001 1.000 1.000 11.895 < 0.001 1.000 1.000 0.000 0.987
Year 2.123 2.580 1.641 0.190 2.723 3.243 3.228 0.024 1.000 1.000 0.022 0.884
Body mass 3.801 3.955 5.689 < 0.001 2.897 3.415 13.967 < 0.001 2.021 2.527 3.934 0.016
Pop. size 2.584 3.069 8.390 < 0.001 1.000 1.000 0.017 0.896 3.972 3.997 2.653 0.039
Carnivores 2.640 3.201 2.898 0.038 1.000 1.000 5.524 0.021 3.518 3.876 8.167 < 0.001
Insectivores 3.148 3.553 3.466 0.016 1.847 2.249 3.268 0.038 2.027 2.470 2.113 0.115
Omnivores 3.936 3.990 8.461 < 0.001 2.503 3.004 1.626 0.189 1.000 1.000 10.444 < 0.01
Granivores 2.927 3.361 4.707 < 0.01 3.527 3.782 4.148 < 0.01 1.000 1.000 9.560 < 0.01
Scavengers 3.836 3.961 5.843 < 0.001 3.506 3.825 6.844 < 0.001 2.368 2.809 2.756 0.051
Land-use*PC1 climate 1.073 1.073 1.893 0.170 1.077 1.077 0.396 0.546 1.077 1.077 0.006 0.947
36
Land-use*PC2 climate 2.098 2.513 0.626 0.573 1.077 1.077 1.728 0.189 1.077 1.077 0.181 0.691
Land-use*Space 2.561 2.974 3.555 0.019 1.422 1.650 1.955 0.149 2.461 2.905 0.969 0.406
Land-use*Area 1.073 1.073 1.829 0.178 1.077 1.077 1.468 0.227 2.275 2.631 2.794 0.052
Land-use*Year 2.164 2.620 1.749 0.168 1.077 1.077 0.761 0.389 1.077 1.077 0.191 0.682
Land-use*Body mass 2.881 3.051 6.926 < 0.001 1.077 1.077 0.064 0.819 1.077 1.077 0.122 0.747
Land-use*Pop. size 1.073 1.073 1.062 0.306 1.077 1.077 0.930 0.339 1.077 1.077 1.371 0.243
Land-use*Carnivores 1.073 1.073 4.491 0.035 1.077 1.077 2.729 0.099 1.077 1.077 0.455 0.516
Land-use*Insectivores 3.971 4.047 7.788 < 0.001 2.854 3.304 0.906 0.446 2.701 3.171 1.853 0.140
Land-use*Omnivores 1.073 1.073 0.936 0.338 1.077 1.077 2.137 0.144 1.077 1.077 0.305 0.560
Land-use*Granivores 1.917 2.293 2.353 0.095 2.732 3.234 1.014 0.392 3.724 3.978 2.916 0.026
Land-use*Scavengers 1.073 1.073 1.761 0.186 1.077 1.077 1.938 0.164 2.010 2.116 0.491 0.624
37
R-sq (adj.) 0.964 0.898 0.704
GSV score 124.560 0.963 0.527
Dev. Explained 98.1% 93.4% 80.4%
38
Figure 7. Scatterplot of the Generalized Additive Model for European-wide assemblage relative to
species richness versus area (log in km2). Red line represents the fit evaluated at the mean of the
independent variable and blue lines represent the fitted lines of ± two times the standard error of the
independent variable.
Relative to the effect of the land-use factor, it was only significant for
species richness regarding species’ body mass (g) and percentage of
Insectivores, indicating significant differences between natural and antrhopogenic
sets for both variables (Table 5). Studies composed of bird species with average
body mass between 50 and 100 g are those who contribute more to species
richness at European-wide scale. It’s in natural sites where this variable reaches
its highest value and also where it can be found the sites with the biggest animals
(Figure 8). Considering the percentage of insectivorous species, it presents an
assymptotic relationship with species richness, with higest richness in natural
sites where there are about 40% insectivores and in anthropogeic sites where
there are about 55% insectivores, assemblage wide. It’s also in natural sites
where it can be found the highest percentage of insectivores (Figure 8).

39
Figure 8. Scatterplots of the Generalized Additive Model for European-wide assemblage relative to species
richness versus body mass (g) (on top) and percentage of insectivores (on bottom), with significant
differences considering land-use. Red line represents the fit evaluated at the mean of both independent
variables. Blue dots represent natural sites; black dots represent antrhopogenic sites.
40
4. Discussion
4.1 Spatial distribution and phylogenetic structure of assemblages
Assemblage-wide analysis of co-occurrence as well as overall patterns
of phylogenetic relatedness within assemblages measured by the mean
phylogenetic distance as well as by the mean nearest neighbour distance
showed that, regardless of considered grain, environmental filtering is more
important than competitive interactions in shaping bird assemblages.
This is similar to the results obtain for bats within Bavaria (Riedinger et
al. 2013) but contradicts the findings from Denmark (Gotelli et al. 2010), which
showed signs of competition across a similar spatial scale. Like Denmark,
present-day Bavaria offers an ideal geographic setting for co-occurrence
analysis of avian species at regional scale, since there are no major geographic
barriers to avian dispersal. Furthermore, no in situ speciation occurred in
Bavaria, otherwise endemic species or at least subspecies should occur.
However, Gotelii et al. (2010) analysed the structure within selected guilds
whereas the present study used all genera observed. Therefore, the present
approach might have diluted signs of competition.
However, using the approach followed on the present study it is possible
to conclude that across all genera which may occur on a grid, competition is in
general not important, given the assumption that the intensity of competitive
interactions decreases with increasing phylogenetic intensity. Therefore, the
present analysis is not showing that competition is not occurring; it states only
that it is not important for the distribution of species at the considered grain.
41
Although it might seem contradictory that similar species can be able to
co-exist and competitive interactions might be overcame, in other studies
(Stamps 1991; Stephens & Sutherland 1999) it has been acknowledge that a
positive relationship between the suitability of a patch and the number of
conspecifics is not uncommon, leading to aggregative behaviour, even in
territorial birds. This, allied with the fact that sometimes bird species can
aggregate to form colonies during breeding season, might contribute to
clustering of the assemblages studied and the co-occurrence found.
4.2 Anthropogenic land-use impact
At the regional level, for Bavaria, and according to previous predictions,
there was an overall decrease of both measures of phylogenetic diversity with
increased percentage of land-use, more specifically, anthropogenic habitats and
meadows. This decrease occurred despite the fact that species richness was
significantly increasing with the percentage of most land-type analysed, even
when only native species were selected.
Both measures of phylogenetic diversity capture a different aspect of the
phylogenetic structure. Mean phylogenetic distance is thought to be more
sensitive to tree-wide patterns of phylogenetic clustering and evenness (Kembel
2010). Mean nearest neighbour distance, which is the mean distance that
separates each species in the community from its closest relative, is thought to
be more sensitive to patterns close to the tips of the phylogeny (Kraft et al.
2007).
This difference is already evident when comparing the distribution of
effect sizes between both measures, since, although both were negative, the
42
effect size of the mean phylogenetic distance was always more negative than
the effect size of the mean nearest neighbour distance.
Considering bird’s evolutionary history, Aves class is divided into
Paleognathae (compromising ratites and tinamous) and Neognathae (all
others); furthermore, the last one is split into Galloanserae (chickens, ducks and
allies) and Neoaves (all others) (Hackett et al. 2008).
The slight, punctual increase of the effect size of mean nearest
neighbour distance with the percentage of anthropogenic habitats (until 15% of
their occupancy) can be explained by the occurrence of numerous species of
birds belonging to the lineage of the Galloanseres. Increasing percentage of
existing water within city parks and gardens within anthropogenic habitats leads
to momentary increase of these species, highly associated with water
availability and/or open habitats.
Furthermore, this lineage diverged from the Neoaves in the Mid-
Cretaceous (120 to 90 M.y. ago). When analysing the effect size of the mean
phylogenetic distance all pairs of species belonging to both lineages are
included and therefore the influenced of Galloanseres presence is deluded,
leading to decreasing values. In contrast, the effect size of the mean nearest
neighbour distance includes only pairs within each lineage and its value
becomes higer. This suggests that habitat affiliation evolved very early in the
history of birds [e.g. Prinzing et al. (2001) for plants].
In agreement with other studies from plants in Germany (Knapp et al.
2008) and bats in Bavaria (Riedinger et al. 2013) in the present study and in
particular for anthropogenic habitats, although species richness increased,
phylogenetic diversity decreased with the percentage of such land-type. This
43
means that, also for birds, the necessary traits to cope with the pressures of
urban habitats are confined to certain lineages, less sensitive to human
presence. It may indicate that with the further increase of species with
increasing amount of urban habitats, species from already occurring lineages
are added to the assemblages, a clear sign of habitat filtering.
Overall, the hump-shaped relationship found between the effect size of
the mean nearest neighbour distance and anthropogenic habitats can be
explained by the intermediate disturbance hypothesis (Wilkinson 1999). A small
level of urbanization within an area can lead to more complex and
heterogeneous conditions, supporting more species of birds, and consequently,
more different lineages, especially in suburban areas at the interface between
urban and more natural areas (Andersson 2006; Blair 2004). This increase in
environmental heterogeneity and resource availability promotes increasing
structural aspects of the environment, allowing more species from different
lineages to be added to the assemblage.
Regarding agriculture, only species richness was related to the
percentage of this land-type. Although non-significant, the relationship between
percentage of agriculture and phylogenetic diversity was positive. This might be
explained by the fact that, although half of Bavarian territory is classified as
agricultural land, according to data from the Bavarian Ministry of Agriculture and
Forestry (http://www.stmelf.bayern.de) most farm holdings are small (less than
50 ha) and those with highest dimensions are under strict management actions.
So, although agriculture is a land-type highly developed within Bavaria, its
impact might be deluded by strong management actions, therefore leading to an
44
increase in species richness and having a positive influence on bird
phylogenetic diversity.
Contrary to agriculture, the effect size of the mean phylogenetic distance
was negatively related to the percentage of meadows. Such land-type can be
defined as grassland not grazed by domestic livestock and, within Bavaria, most
agricultural land is occupied by crop land, especially cereal yield. Therefore
such land-type leads to a decrease in bird phylogenetic diversity, through
habitat filtering.
At continental-wide scale sample area seems to be the most important
factor shaping species richness and mean phylogenetic distance, with
increasing number of species and mean phylogenetic distance with increasing
area (km2). This is explained accordingly to the More Individuals Hypothesis
(MIH), which predicts a positive relationship between species richness and
available energy. Such hypothesis has been suggested for bird communities
(Hurlbert 2004), assuming that larger areas contain greater food resources,
therefore supporting more individuals; communities with more individuals
include also different species. A larger number of species distantly related leads
to an increase of mean phylogenetic distance with increasing area.
However, this also contradicts previous statements (Vamosi et al. 2009),
since competitive interactions were thought to be more important at small
spatial scales whereas habitat filtering should became more important at larger
scales. In the present study, as the study-area increases the effect size of the
mean phylogenetic distance assumes higher values, with more unrelated
species being added to the assemblage. This indicates phylogenetic
overdispersion of assemblages for bigger areas. Since overdispersion means
45
that less related species are co-occurring, is normally caused by competitive
interactions. Therefore, at smaller spatial scales habitat filtering seems to be
more important in shaping European-wide bird assemblage, whereas at bigger
spatial scales, competitive interactions have a major role. This might be
explained by the fact that, as the area increases more species, and more
lineages, are added to the assemblage. Species closely related will compete for
resources and eliminate each other, ultimately leading to an assemblage where
the mean distance that separates each species from the tree root – the mean
phylogenetic distance– assumes higher values, and the effect size assumes
positive ones.
Climate, summarized by PC1, also had a positive relationship with
species richness and mean nearest neighbour distance, with increasing number
of species and phylogenetic diversity with increasing temperature. This is
explained according to the metabolic theory, which states that almost all rates of
biological acitivity increase with temperature, leading to increasing energy and
productivity (Brown et al. 2004). According to Allen et al. (2007) community
species abundance is predicted to increase with the rate of consumptition of net
primary production, which in turn, increases with increasing temperature.
Therefore, available ecosystem energy helps to accumulate biodiversity through
an increase in species richness and also increasing phylogenetic diversity.
Time had no significant effect upon any response variable analysed
within Europe. Since only thirty-five years were represented in the present
study, it is thought to be not enough time to show a response upon richness or
phylogeny of the assemblage. However non-significant it was registered an
increase of all biodiversity variables with time; this is overall optimistic, since it
46
confirms that the management actions performed within European Union to
improve biodiversity, especially regarding agriculture, are presenting good
results (EEA 2010).
Contrary to previous predictions, there were no significant overall
differences between natural and antrhopogenic sites within Europe. Only two
trait variables analysed at the species richness level where significantly different
between land-use: specie’s body mass (g) and percentage of insectivores.
In natural sites one can find the biggest animals; this is explained by the
fact that bigger animals need bigger home ranges, with heterogeneity of offered
resources. In a study about the body mass of house sparrows (Passer
domesticus), one of the most common species recorded within urban areas,
Liker et al. (2008) discovered that birds within more urbanized areas were
smaller than the ones living in natural areas and such difference was probably
due to adaptative divergence to environmental variances. In the present study
this is confirmed for all species dated within Europe.
Urban areas, as mentioned before, benefit from highly favourable climatic
conditions, which may contribute to an increase in insect biomass, leading to
changes in urban bird communities at the throphic level (Faeth et al. 2005), with
bird species feeding primarily on this higly available, energetic food resource,
especially during breeding season. This leads to a maximum of urban species
richness when there is about 55% insectivores within the assemblage, contrary
to the 40% registered within natural sites.
47
4.3 Implications for conservation
The present study holds important results regarding human impact upon
the structure of bird assemblages. Its results focus on urban impact, considered
the most anthropogenically transformed, permanent and fastest growing habitat
in the world. As proved by this study, although its heterogeneity might provide
food resources availability, and consequently, more niches (Stevens et al.
2011), leading to a surprising increase in species richness, this is only true for
certain species, both at regional and continental scale (Chace & Walsh 2006).
Urban assemblages are composed of a few, certain lineages that have
specific traits that allow them to cope with this particular environment.
Therefore, increasing urban spread throughout the world could untimely lead to
a phenomenon of biotic homogenization (Gregory et al. 2005; Olden 2006),
where assemblages become more and more phylogenetically alike, with similar
lineages and species.
Anthropogenic habitats can, in this way, be a potent selective force
capable of shaping the behaviour of species [e.g. voice characteristics
(Slabbenkoorn & Peet 2003) and flight distance (Moller 2008)] and interactions
between organisms (Moller 2009). This may lead, in the longer term, to change
in population genetics acting also as a powerful evolutionary force (Grimm et al.
2008).
It is important to perform more phylogenetic assemblage studies similar
to this one, particularly comparing several metrics and scales, to understand the
processes that influence assemblages under growing anthropogenic impact.
The present study showed that regionally, although species richness might be
favoured by increasing antrhopogenic habitats, the same is not true for
48
phylogenetic diversity. Also, farmland seems to be the habitat with the least
impact upon bird assemblages within Bavaria. At a continental scale, for
Europe, it was showen that bot spatial area and climate have a positive
influence upon both species richness and phylogenetic diversity, pressing the
issue towards a better management, with more continuous areas of
heterogeneous habitat, especially in southern Europe, where climate is more
favourable, particularly during breeding season.
Together with experimental studies [e.g. Tilman et al. (2001) and Spehn
et al. (2005) on plants; Hairston Jr. et al. (2005) on birds; and Ezard et al.
(2009) on mammals], it might be possible in the future to complement
conservation biology with urbanization and agriculture. With proper
management planning future anthropogenic land-use might have the right
characteristics and dimensions that allow them to aggregate large and
phylogenetically heterogeneous communities, contributing to worldwide
ecological diversity.
49
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6. Appendix
Appendix A. Map retrieved from Bezzel et al. (2005) representing an example of a species distribution
map used to collect the regional data in the present study. Here all 2285 grid cells are represented, but
only those who where fully within Bavaria – 1927 - where used for the analysis.
59
Appendix B. List of the 157 bird species recorded in Bavaria between 1996 and 1999 by alphabetic order of the species name. Species are
presented by their scientific name. Nº observations is the value of observed presences for the sum of all grid cells during the same period. Status
represents the status of the species according to the UICN Red List of Threated Species (http://www.iucnredlist.org/); LC – Least Concern; NE –
Near Threatened. Trend stands for population trend in Germany at present days according to accounts in Birds in Europe: population estimates,
trends and conservation status (Birdlife International 2004 – http://www.birdlife.org). Near threatened and threatened species are highlighted.
Species Nº observations Order Family Status Trend
Accipiter gentilis 1107 Falconiformes Accipitridae LC Secure

Accipiter nisus 1298 Falconiformes Accipitridae LC Secure
Acrocephalus arundinaceus 102 Passeriformes Sylviidae LC Secure
Acrocephalus palustris 1556 Passeriformes Sylviidae LC Secure
Acrocephalus schoenobaenus 99 Passeriformes Sylviidae LC Secure
Acrocephalus scirpaceus 841 Passeriformes Sylviidae LC Secure
Actitis hypoleucos 179 Charadriiformes Scolopacidae LC Declining
Aegithalos caudatus 985 Passeriformes Aegithalidae LC Secure
Aegolius funereus 355 Strigiformes Strigidae LC Secure
Alauda arvensis 1745 Passeriformes Alaudidae LC Depleted
Alcedo atthis 719 Coraciiformes Alcedinidae LC Depleted
Anas crecca 187 Anseriformes Anatidae LC Secure
Anas platyrhynchos 1773 Anseriformes Anatidae LC Secure
Anas strepera 139 Anseriformes Anatidae LC Depleted
Anser anser 150 Anseriformes Anatidae LC Secure
60
Anthus pratensis 355 Passeriformes Motacillidae LC Secure

Anthus trivialis 1372 Passeriformes Motacillidae LC Secure
Apus apus 1435 Apodiformes Apodidae LC Secure
Ardea cinerea 618 Ciconiiformes Ardeidae LC Secure
Asio otus 924 Strigiformes Strigidae LC Secure
Aythya ferina 298 Anseriformes Anatidae LC Declining
Aythya fuligula 1017 Anseriformes Anatidae LC Declining
Bonasa bonasia 119 Galliformes Phasianidae LC Secure
Bubo bubo 204 Strigiformes Strigidae LC Depleted
Buteo buteo 1855 Falconiformes Accipitridae LC Increasing
Carduelis cannabina 1146 Passeriformes Fringillidae LC Declining
Carduelis carduelis 1745 Passeriformes Fringillidae LC Secure
Carduelis chloris 1871 Passeriformes Fringillidae LC Secure
Carduelis flammea 294 Passeriformes Fringillidae LC Decreasing
Carduelis spinus 687 Passeriformes Fringillidae LC Secure
Certhia brachydactyla 1342 Passeriformes Certhiidae LC Secure
Certhia familiaris 1378 Passeriformes Certhiidae LC Stable
Charadrius dubius 435 Charadriiformes Charadriidae LC Secure
Ciconia ciconia 154 Ciconiiformes Ciconiidae LC Depleted
Ciconia nigra 126 Ciconiiformes Ciconiidae LC Rare
61
Cinclus cinclus 735 Passeriformes Cinclidae LC Secure

Circus aeruginosus 354 Falconiformes Accipitridae LC Secure
Coccothraustes coccothraustes 1135 Passeriformes Fringillidae LC Secure
Columba oenas 679 Columbiformes Columbidae LC Secure
Columba palumbus 1876 Columbiformes Columbidae LC Secure
Corvus corax 399 Passeriformes Corvidae LC Secure
Corvus corone 1841 Passeriformes Corvidae LC Secure
Corvus monedula 530 Passeriformes Corvidae LC Secure
Coturnix coturnix 807 Galliformes Phasianidae LC Depleted
Crex crex 118 Gruiformes Rallidae LC Depleted
Cuculus canorus 1656 Cuculiformes Cuculidae LC Secure
Cygnus olor 623 Anseriformes Anatidae LC Secure
Delichon urbicum 1823 Passeriformes Hirundinidae LC Declining
Dendrocopos major 1879 Piciformes Picidae LC Secure
Dendrocopos medius 281 Piciformes Picidae LC Secure
Dendrocopos minor 570 Piciformes Picidae LC Secure
Dryocopus martius 1423 Piciformes Picidae LC Secure
Emberiza citrinella 1811 Passeriformes Emberizidae LC Secure
Emberiza schoeniclus 1071 Passeriformes Emberizidae LC Secure
Erithacus rubecula 1902 Passeriformes Muscicapidae LC Secure
62
Falco peregrinus 101 Falconiformes Falconidae LC Secure

Falco subbuteo 708 Falconiformes Falconidae LC Secure
Falco tinnunculus 1749 Falconiformes Falconidae LC Declining
Ficedula albicollis 160 Passeriformes Muscicapidae LC Secure
Ficedula hypoleuca 642 Passeriformes Muscicapidae LC Secure
Fringilla coelebs 1916 Passeriformes Fringillidae LC Secure
Fulica atra 1220 Gruiformes Rallidae LC Secure
Gallinago gallinago 317 Charadriiformes Scolopacidae LC Declining
Gallinula chloropus 1080 Gruiformes Rallidae LC Secure
Garrulus glandarius 1850 Passeriformes Corvidae LC Secure
Glaucidium passerinum 343 Strigiformes Strigidae LC Secure
Hippolais icterina 1189 Passeriformes Sylviidae LC Secure
Hirundo rustica 1864 Passeriformes Hirundinidae LC Depleted
Jynx torquilla 402 Piciformes Picidae LC Declining
Lanius collurio 1544 Passeriformes Laniidae LC Depleted
Larus ridibundus 187 Charadriiformes Laridae LC Secure
Locustella fluviatilis 180 Passeriformes Sylviidae LC Secure
Locustella naevia 1021 Passeriformes Sylviidae LC Secure
Loxia curvirostra 707 Passeriformes Fringillidae LC Secure
Lullula arborea 148 Passeriformes Alaudidae LC Depleted
63
Luscinia megarhynchos 315 Passeriformes Muscicapidae LC Secure

Luscinia svecica 289 Passeriformes Muscicapidae LC Secure
Mergus merganser 211 Anseriformes Anatidae LC Secure
Miliaria calandra 153 Passeriformes Emberizidae LC Declining
Milvus migrans 343 Falconiformes Accipitridae LC Unkown
Milvus milvus 482 Falconiformes Accipitridae NT Declining
Motacilla alba 1895 Passeriformes Motacillidae LC Secure
Motacilla cinerea 1410 Passeriformes Motacillidae LC Secure
Motacilla flava 683 Passeriformes Motacillidae LC Secure
Muscicapa striata 1473 Passeriformes Muscicapidae LC Depleted
Nucifraga caryocatactes 542 Passeriformes Corvidae LC Secure
Numenius arquata 165 Charadriiformes Scolopacidae NT Declining
Oriolus oriolus 716 Passeriformes Oriolidae LC Secure
Parus ater 1751 Passeriformes Paridae LC Stable
Parus caeruleus 1894 Passeriformes Paridae LC Secure
Parus cristatus 1446 Passeriformes Paridae LC Declining
Parus major 1918 Passeriformes Paridae LC Increasing
Parus montanus 1220 Passeriformes Paridae LC Secure
Parus palustris 1472 Passeriformes Paridae LC Decreasing
Passer domesticus 1857 Passeriformes Passeridae LC Declining
64
Passer montanus 1712 Passeriformes Passeridae LC Declining

Perdix perdix 882 Galliformes Phasianidae LC Vulnerable
Pernis apivorus 488 Falconiformes Accipitridae LC Stable
Phoenicurus ochruros 1885 Passeriformes Muscicapidae LC Secure
Phoenicurus phoenicurus 910 Passeriformes Muscicapidae LC Depleted
Phylloscopus bonelli 106 Passeriformes Sylviidae LC Declining
Phylloscopus collybita 1910 Passeriformes Sylviidae LC Secure
Phylloscopus sibilatrix 1360 Passeriformes Sylviidae LC Declining
Phylloscopus trochilus 1829 Passeriformes Sylviidae LC Secure
Pica pica 1687 Passeriformes Corvidae LC Secure
Picus canus 650 Piciformes Picidae LC Depleted
Picus viridis 1182 Piciformes Picidae LC Stable
Podiceps cristatus 413 Podicipediformes Podicipedidae LC Secure
Prunella modularis 1813 Passeriformes Prunellidae LC Secure
Pyrrhula pyrrhula 1505 Passeriformes Fringillidae LC Secure
Rallus aquaticus 259 Gruiformes Rallidae LC Secure
Regulus ignicapilla 1647 Passeriformes Reguliidae LC Stable
Regulus regulus 1684 Passeriformes Reguliidae LC Decreasing
Remiz pendulinus 157 Passeriformes Remizidae LC Increasing
Riparia riparia 282 Passeriformes Hirundinidae LC Depleted
65
Saxicola rubetra 539 Passeriformes Muscicapidae LC Secure

Scolopax rusticola 487 Charadriiformes Scolopacidae LC Declining
Serinus serinus 1594 Passeriformes Fringillidae LC Secure
Sitta europaea 1848 Passeriformes Sittidae LC Secure
Streptopelia decaocto 1666 Columbiformes Columbidae LC Secure
Streptopelia turtur 607 Columbiformes Columbidae LC Declining
Strix aluco 1330 Strigiformes Strigidae LC Secure
Sturnus vulgaris 1871 Passeriformes Sturnidae LC Declining
Sylvia atricapilla 1899 Passeriformes Sylviidae LC Secure
Sylvia borin 1722 Passeriformes Sylviidae LC Secure
Sylvia communis 1130 Passeriformes Sylviidae LC Secure
Sylvia curruca 1345 Passeriformes Sylviidae LC Secure
Tachybaptus ruficollis 732 Podicipediformes Podicipedidae LC Secure
Tetrao urogallus 113 Galliformes Phasianidae LC Secure
Troglodytes troglodytes 1896 Passeriformes Troglodytidae LC Secure
Turdus merula 1918 Passeriformes Turdidae LC Secure
Turdus philomelos 1899 Passeriformes Turdidae LC Secure
Turdus pilaris 1789 Passeriformes Turdidae LC Secure
Turdus torquatus 110 Passeriformes Turdidae LC Secure
Turdus viscivorus 1594 Passeriformes Turdidae LC Secure
66
Tyto alba 400 Strigiformes Tytonidae LC Declining

Vanellus vanellus 952 Charadriiformes Charadriidae LC Vulnerable
67
Appendix C. List of the 127 studies that made out the European-wide analysis, listed by alphabetic
order of the country, and within country by name of the place/city of each study. It is also attached the
correspondent list of references.
Country Place/City
Dürrenstein (Leditznig & Pekny 2008)
Krappfeld Kärten (Lentner 1997)
Austria Tirol (Landmann 1996)
Vienna (Sziemer & Holzer 2005)
Vienna (Wichmann & Purtscher 2009)
Belgium Brussel (Weiserbs & Jacob 2005)
Bulgaria Sofia (Iankov 2005)
Czech Republic Prague (Stastný et al. 2005)
Seskar Arquipelago (Vasilyeva 2002)
Finland Suurpelto Agricultural Area (Heikkinen & Korpela 2001)
Tornio (Huhtalo & Järvinen 1977)
Caen (Lang 2006)
Grand Haze Marshes (Lecocq 1992)
Le Havre (Lang 2006)
Montepellier (Caula et al. 2008)
France
Normandie (Normand 1992)
Rouen (Lang 2006)
Saint-Lo (Lang 2006)
Saint-Severe Forerst (Bruno 2005)
Aberseewand (Scherzinger 1982)
Auer Weidmoos Rosenheim (Nitsche 2004)
Augsburg (Bauer 2000)
Bayern (Bezzel et al. 2005)
Berlin (Witt 2005)
Bielefeld (Laske 1991)
Germany
Bodensee (Bauer & Heine 1992)
Bonn (Rheinwald 2005)
Bremen (Seitz & Dallmann 1992)
Chemnitz (Flöter et al. 2006)
Fränkischen Weihergebiet (Kraus & Krauss 2003)
Grossraum Bonn (Wink 1980)
68
Country Place/City
Hagen (Welzel 2009)
Haidenaabtal/Oberpfalz (Bastian 1993)
Hamburg (Musow 2005)
Hoyerswerda-Neustadt (Krüger 1973)
Kemnather Hügelland (Möhrlein 2001a)
Kreis & Stadt Ansbach (Ranftl & Dornberger 2002)
Kreis und Stadt Würzburg (Uhlich 1991)
Kreises Soest (Illner et al. 1989)
Kreises Waren (Kremp & Krägenow 1976)
Landkreis Eichsfeld (Hartmann 2004)
Mainz (Thomas 1983)
Mönchengladbach (Hurtmann 2005)
Germany Osnabrück (Kooiker 1994)
Ostdeutschlad (Nicolai 1993)
Ostoberfranken (Gubitz et al. 1993)
Plössberger Hügelland (Möhrlein 2001b)
Rheinland (Nordrhein) (Wink et al. 2005)
Rötelseeweihergebietes (Zach 2002)
Rückhaltebeckens Straussfurt (Laussmann & Frick 2008)
Sachsen (Steffens et al. 1998)
Stadtgebiet Eisenach (Mey 2005)
Stadtgebiet Nürnberg (Veitengruber 1995)
Thüringen (Rost & Grimm 2004)
Wasserburg & Rosenheim (Mieslinger 1997)
Werdenfelser Land (Bezzel & Lechner 1978)
Alta Valsessera (Popy et al. 2010)
Bergamo (Cairo et al. 2006)
Campagnia (Atripaldi et al. 1989)
Carpeneto County (Spano 1984)
Distritto Mendrisiotto (Lardelli 1988)
Italy
Florence (Dinetti 2005)
Genova (Borgo et al. 2005)
Monte Goadagnolo (Lorenzetti et al. 2004)
Monti Simbruini Regional Park (de Pisi & Fusacchia 2005)
Piemonte e Val d'Aosta (Mingozzi et al. 1988)
69
Country Place/City
Provincia Verona (de Franceschi 1991)
River Serchio (Tuscany) (Verducci & Chines 2009)
Italy
Rome (Cignini & Zapparoli 2005)
Torino (Maffei 2001)
Lithuania Lake Kretuonas (Logminas & Rianba 1999)
Moldova Landscape Park Izmailskie Islands (Potapov 2001)
Bagna Struskie Bogs (Solowej & Wysocki 2001)
Bialowieza National Park (Wesolowski et al. 2003)
Bystrzyckie Mountains (Mikusek 1996)
Damnica (Gorski 1988)
Former Military Camp North Przemkow (Adamski & Czapulak 2002)
Gliwic (Betleja 2007)
Krkonose Biosphere Reserve (Flousek & Gramsz 1999)
Lake Luknajno (Osojca 2005)
Lower Narew Valley (Rzepala et al. 1999)
Lublin (Biadun 2005)
Meadows Lake Miedwie (Guentzel & Wysocki 2004)
Miedzyodrze Area (Lawicki et al. 2007)
Modrzewina Natural Reserve (Chmielewski 1992)
Ner River Valley (Mielczarek 2006)
Poland
Nida River Valley (Polak & Wilniewczyc 2001)
Odra Valley (Hedba & Wyszynski 2002)
Ogrodniki (Golawski & Dombrowski 2004)
Olszyn (Novakowski 1996)
Paprotina (Golawski & Dombrowski 2004)
Pilica River Floodland (Chmielewski et al. 1993)
Potegowo (Gorski 1988)
Protection Forest Szast (Zmihorski 2008)
Przemysl province (Hordowski & Kunysz 1991)
Stolowe Mountains (Mikusek & Dyrcz 2003)
Tarnow region (Martyka et al. 2002)
Tomaszowa Mazowieckiego (Sosnowski 1994)
Torun (Zalewski 1994)
Varsaw (Luniak 2005)
70
Country Place/City
Lisbon (Geraldes & Costa 2005)
Minho (Moreira et al. 2001)
Portugal
Paul do Taipal (Tenreiro 2002)
Serra da Nogueira (Patacho 2002)
Moscow (Konstantinov & Zakharov 2005)
Russia
St. Petersburg (Khrabryi 2005)
Bratislava (Feriancová-Masárová & Kalivodová 2005)
Gory Opawskie Landscape Park (Hedba 2001)
Slovakia
Prievidza (Sotnár 1994)
Senne Fishponds (Wieland 1999)
Slovenia Drawa River (Braèko 1997)
Alto Vinalopó (Alicante) (Campos 2001)
Catalunya (Llobet & Estrada 2004)
Comunidad de Madrid (Martí 1994)
Comunidad de Valencia (Polo & Polo 2003)
Spain Comunidad Valenciana (Moliner 1991)
Menorca (Salom 1997)
Navarra (Aldaroso 1985)
Parque Nacional Aigüestortes (Blanch 2005)
Valencia (Murgui 2005)
Sweden Örebro (Sandström & Mikusinski 2006)
Ukraine Bozhanovo (Lukashuk 1996)
Cheshire and Wirral (Norman 2008)
County Durham (Bowey & Westerberg 2000)
United Kingdom Lundy (Davis et al. 2007)
Malvern Hills (Duncan 2008)
Sheffield (Fuller et al. 2009)
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80
Appendix D. List of the 297 bird species recorded for European-wide analysis by alphabetic order of the species name. Species are presented
by their scientific name. Nº observ. Natural is the value of observed presences for the sum of all studies within natural sites and Nº observ.
Anthropogenic is the value of observed presences for the sum of all studies within anthropogenic sites. Status represents the status of the
species according to the UICN Red List of Threated Species (http://www.iucnredlist.org/); LC – Least Concern; NE – Near Threatened. Trend
stands for population trend in Europe at present days according to accounts in Birds in Europe: population estimates, trends and conservation
status (Birdlife International 2004 – http://www.birdlife.org). Near Threatened and Threatened species are highlighted.
Nº observ. Nº observ.
Species Order Family Status Trend
Natural Urban
Accipiter gentilis 54 24 Falconiformes Accipitridae LC Secure
Accipiter nisus 57 30 Falconiformes Accipitridae LC Secure
Acrocephalus arundinaceus 41 16 Passeriformes Sylviidae LC Secure
Acrocephalus palustris 44 28 Passeriformes Sylviidae LC Secure
Acrocephalus schoenobaenus 36 17 Passeriformes Sylviidae LC Secure
Acrocephalus scirpaceus 52 28 Passeriformes Sylviidae LC Secure
Actitis hypoleucos 34 12 Charadriiformes Scolopacidae LC Declining
Aegithalos caudatus 64 37 Passeriformes Aegithalidae LC Secure
Aegolius funereus 20 2 Strigiformes Strigidae LC Secure
Aegypius monachus 1 0 Falconiformes Accipitridae NT Rare
Alauda arvensis 63 36 Passeriformes Alaudidae LC Depleted
Alca torda 2 0 Charadriiformes Alcidae LC Secure
Alcedo atthis 55 26 Coraciiformes Alcedinidae LC Depleted
81
Natural Urban
Alectoris rufa 13 2 Galiiformes Phasianidae LC Declining
Anas acuta 8 2 Anseriformes Anatidae LC Declining
Anas clypeata 34 8 Anseriformes Anatidae LC Declining
Anas crecca 32 11 Anseriformes Anatidae LC Secure
Anas penelope 9 1 Anseriformes Anatidae LC Secure
Anas platyrhynchos 68 39 Anseriformes Anatidae LC Secure
Anas querquedula 36 11 Anseriformes Anatidae LC Declining
Anas strepera 31 8 Anseriformes Anatidae LC Depleted
Anser anser 25 9 Anseriformes Anatidae LC Secure
Anthus campestris 26 6 Passeriformes Motacillidae LC Declining
Anthus pratensis 44 22 Passeriformes Motacillidae LC Secure
Anthus spinoletta 17 2 Passeriformes Motacillidae LC Secure
Anthus trivialis 59 28 Passeriformes Motacillidae LC Secure
Apus apus 53 43 Apodiformes Apodidae LC Secure
Apus pallidus 6 6 Apodiformes Apodidae LC Secure
Aquila chrysaetos 17 1 Falconiformes Accipitridae LC Rare
Aquila fasciatus 6 1 Falconiformes Accipitridae LC Endangered
Aquila pomarina 11 0 Falconiformes Accipitridae LC Declining
Ardea cinerea 38 12 Ciconiiformes Ardeidae LC Secure
Ardea purpurea 14 2 Ciconiiformes Ardeidae LC Declining
Ardeola ralloides 5 1 Ciconiiformes Ardeidae LC Declining
82
Natural Urban
Arenaria interpres 1 0 Charadriiformes Scolopacidae LC Secure
Asio flammeus 9 4 Strigiformes Strigidae LC Depleted
Asio otus 60 27 Strigiformes Strigidae LC Secure
Athene noctua 39 22 Strigiformes Strigidae LC Declining
Aythya ferina 32 10 Anseriformes Anatidae LC Declining
Aythya fuligula 42 22 Anseriformes Anatidae LC Declining
Aythya nyroca 11 2 Anseriformes Anatidae NT Vulnerable
Bombycilla garrulus 2 0 Passeriformes Bombycillidae LC Secure
Bonasa bonasia 15 2 Galiiformes Phasianidae LC Secure
Botaurus stellaris 21 5 Ciconiiformes Ardeidae LC Depleted
Branta leucopsis 4 0 Anseriformes Anatidae LC Secure
Bubo bubo 31 6 Strigiformes Strigidae LC Depleted
Bubulcus ibis 5 1 Ciconiiformes Ardeidae LC Secure
Bucephala clangula 11 6 Anseriformes Anatidae LC Secure
Burhinus oedicnemus 11 1 Charadriiformes Burhinidae LC Vulnerable
Buteo buteo 69 30 Falconiformes Accipitridae LC Increasing
Calandrella brachydactyla 10 2 Passeriformes Alaudidae LC Declining
Calidris alpina 8 0 Charadriiformes Scolopacidae LC Depleted
Calonectris diomedea 4 0 Procellariiformes Procellariidae LC Vulnerable
Caprimulgus europaeus 38 10 Caprimulgiformes Caprimulgidae LC Depleted
Caprimulgus ruficollis 6 1 Caprimulgiformes Caprimulgidae LC Secure
83
Natural Urban
Carduelis cannabina 62 35 Passeriformes Fringillidae LC Declining
Carduelis carduelis 66 41 Passeriformes Fringillidae LC Secure
Carduelis chloris 70 47 Passeriformes Fringillidae LC Secure
Carduelis citrinella 9 0 Passeriformes Fringillidae LC Secure
Carduelis flammea 28 10 Passeriformes Fringillidae LC Decreasing
Carduelis spinus 40 15 Passeriformes Fringillidae LC Secure
Carpodacus erythrinus 21 9 Passeriformes Fringillidae LC Secure
Casmerodius albus 5 1 Ciconiiformes Ardeidae LC Unkown
Certhia brachydactyla 57 32 Passeriformes Certhiidae LC Secure
Certhia familiaris 50 19 Passeriformes Certhiidae LC Stable
Cettia cetti 16 7 Passeriformes Sylviidae LC Secure
Charadrius alexandrinus 8 0 Charadriiformes Charadriidae LC Declining
Charadrius dubius 50 29 Charadriiformes Charadriidae LC Secure
Charadrius hiaticula 11 5 Charadriiformes Charadriidae LC Secure
Chen caerulescens 1 0 Anseriformes Anatidae LC Secure
Chersophilus duponti 2 0 Passeriformes Alaudidae NT Depleted
Chlidonias hybrida 7 0 Charadriiformes Laridae LC Depleted
Chlidonias niger 13 5 Charadriiformes Laridae LC Depleted
Ciconia ciconia 38 10 Ciconiiformes Ciconiidae LC Depleted
Ciconia nigra 20 2 Ciconiiformes Ciconiidae LC Rare
Cinclus cinclus 41 13 Passeriformes Cinclidae LC Secure
84
Natural Urban
Circaetus gallicus 12 1 Falconiformes Accipitridae LC Stable
Circus aeruginosus 38 12 Falconiformes Accipitridae LC Secure
Circus cyaneus 13 0 Falconiformes Accipitridae LC Depleted
Circus pygargus 30 3 Falconiformes Accipitridae LC Secure
Cisticola juncidis 14 5 Passeriformes Cisticolidae LC Secure
Clamator glandarius 7 0 Cuculiformes Cuculidae LC Secure
Clangula hyemalis 1 0 Anseriformes Anatidae VU Secure
Coccothraustes coccothraustes 52 29 Passeriformes Fringillidae LC Secure
Columba livia 29 39 Columbiformes Columbidae LC Secure
Columba oenas 44 21 Columbiformes Columbidae LC Secure
Columba palumbus 71 40 Columbiformes Columbidae LC Secure
Coracias garrulus 12 0 Coraciiformes Coraciidae NT Vulnerable
Corvus corax 49 11 Passeriformes Corvidae LC Secure
Corvus corone 66 40 Passeriformes Corvidae LC Secure
Corvus frugilegus 22 21 Passeriformes Corvidae LC Secure
Corvus monedula 52 38 Passeriformes Corvidae LC Secure
Coturnix coturnix 63 23 Galiiformes Phasianidae LC Depleted
Crex crex 38 16 Gruiformes Rallidae LC Depleted
Cuculus canorus 75 33 Cuculiformes Cuculidae LC Secure
Cyanopica cyanus 1 0 Passeriformes Corvidae LC Secure
Cygnus olor 43 23 Anseriformes Anatidae LC Secure
85
Natural Urban
Delichon urbicum 58 42 Passeriformes Hirundinidae LC Declining
Dendrocopos leucotos 9 5 Piciformes Picidae LC Secure
Dendrocopos major 69 39 Piciformes Picidae LC Secure
Dendrocopos medius 29 14 Piciformes Picidae LC Secure
Dendrocopos minor 54 28 Piciformes Picidae LC Secure
Dendrocopos syriacus 3 9 Piciformes Picidae LC Secure
Dryocopus martius 49 24 Piciformes Picidae LC Secure
Egretta garzetta 11 2 Ciconiiformes Ardeidae LC Increasing
Elanus caeruleus 1 0 Falconiformes Accipitridae LC Rare
Emberiza cia 21 3 Passeriformes Emberizidae LC Depleted
Emberiza cirlus 19 6 Passeriformes Emberizidae LC Secure
Emberiza citrinella 59 33 Passeriformes Emberizidae LC Secure
Emberiza hortulana 30 12 Passeriformes Emberizidae LC Depleted
Emberiza schoeniclus 51 29 Passeriformes Emberizidae LC Secure
Eremophila alpestris 2 0 Passeriformes Alaudidae LC Secure
Erithacus rubecula 72 40 Passeriformes Muscicapidae LC Secure
Erythropygia galactotes 3 0 Passeriformes Muscicapidae LC Vulnerable
Eudromias morinellus 3 0 Charadriiformes Charadriidae LC Secure
Falco naumanni 7 0 Falconiformes Falconidae LC Depleted
Falco peregrinus 30 14 Falconiformes Falconidae LC Secure
Falco subbuteo 50 19 Falconiformes Falconidae LC Secure
86
Natural Urban
Falco tinnunculus 62 40 Falconiformes Falconidae LC Declining
Ficedula albicollis 17 7 Passeriformes Muscicapidae LC Secure
Ficedula hypoleuca 44 27 Passeriformes Muscicapidae LC Secure
Ficedula parva 20 12 Passeriformes Muscicapidae LC Stable
Fratercula arctica 2 0 Charadriiformes Alcidae LC Depleted
Fringilla coelebs 75 46 Passeriformes Fringillidae LC Secure
Fulica atra 55 31 Gruiformes Rallidae LC Secure
Fulmarus glacialis 3 0 Procellariiformes Procellariidae LC Secure
Galerida cristata 25 23 Passeriformes Alaudidae LC Depleted
Galerida theklae 8 0 Passeriformes Alaudidae LC Depleted
Gallinago gallinago 39 16 Charadriiformes Scolopacidae LC Declining
Gallinula chloropus 60 37 Gruiformes Rallidae LC Secure
Garrulus glandarius 69 40 Passeriformes Corvidae LC Secure
Glareola pratincola 3 0 Charadriiformes Glareolidae LC Declining
Glaucidium passerinum 19 3 Strigiformes Strigidae LC Secure
Grus grus 14 2 Gruiformes Gruidae LC Depleted
Gypaetus barbatus 3 0 Falconiformes Accipitridae LC Vulnerable
Gyps fulvus 7 0 Falconiformes Accipitridae LC Secure
Haematopus ostralegus 11 2 Charadriiformes Haematopodidae LC Secure
Haliaeetus albicilla 13 2 Falconiformes Falconidae LC Rare
Hieraaetus pennatus 10 0 Falconiformes Falconidae LC Rare
87
Natural Urban
Himantopus himantopus 13 2 Charadriiformes Recurvirostridae LC Secure
Hippolais icterina 42 28 Passeriformes Sylviidae LC Secure
Hippolais polyglotta 18 7 Passeriformes Sylviidae LC Secure
Hirundo daurica 8 1 Passeriformes Hirundinidae LC Increasing
Hirundo rupestris 18 4 Passeriformes Hirundinidae LC Secure
Hirundo rustica 59 41 Passeriformes Hirundinidae LC Depleted
Hydrobates pelagicus 5 0 Procellariiformes Hydrobatidae LC Secure
Ixobrychus minutus 27 13 Ciconiiformes Ardeidae LC Depleted
Jynx torquilla 55 25 Piciformes Picidae LC Declining
Lagopus muta 10 0 Galiiformes Phasianidae LC Secure
Lanius collurio 62 29 Passeriformes Laniidae LC Depleted
Lanius excubitor 39 9 Passeriformes Laniidae LC Depleted
Lanius minor 9 3 Passeriformes Laniidae LC Declining
Lanius senator 17 3 Passeriformes Laniidae LC Declining
Larus argentatus 13 7 Charadriiformes Laridae LC Secure
Larus cachinnans 7 6 Charadriiformes Laridae LC
Larus canus 16 5 Charadriiformes Laridae LC Depleted
Larus fuscus 12 2 Charadriiformes Laridae LC Secure
Larus marinus 6 1 Charadriiformes Laridae LC Secure
Larus melanocephalus 15 3 Charadriiformes Laridae LC Secure
Larus michahellis 6 2 Charadriiformes Laridae LC
88
Natural Urban
Larus ridibundus 28 12 Charadriiformes Laridae LC Secure
Limicola falcinellus 1 0 Charadriiformes Scolopacidae LC Declining
Limosa limosa 22 5 Charadriiformes Scolopacidae NT Vulnerable
Locustella fluviatilis 30 12 Passeriformes Sylviidae LC Secure
Locustella luscinioides 37 10 Passeriformes Sylviidae LC Secure
Locustella naevia 47 21 Passeriformes Sylviidae LC Secure
Loxia curvirostra 41 10 Passeriformes Fringillidae LC Secure
Lullula arborea 45 14 Passeriformes Alaudidae LC Depleted
Luscinia luscinia 18 13 Passeriformes Muscicapidae LC Secure
Luscinia megarhynchos 44 27 Passeriformes Muscicapidae LC Secure
Luscinia svecica 29 8 Passeriformes Muscicapidae LC Secure
Lymnocryptes minimus 2 0 Charadriiformes Scolopacidae LC Declining
Marmaronetta angustirostris 2 0 Anseriformes Anatidae VU Vulnerable
Melanitta fusca 3 0 Anseriformes Anatidae EN
Melanocorypha calandra 7 0 Passeriformes Alaudidae LC Declining
Mergellus albellus 2 0 Anseriformes Anatidae LC Declining
Mergus merganser 15 3 Anseriformes Anatidae LC Secure
Merops apiaster 21 9 Coraciiformes Meropidae LC Depleted
Miliaria calandra 48 21 Passeriformes Emberizidae LC Declining
Milvus migrans 34 7 Falconiformes Accipitridae LC Unkown
Milvus milvus 30 9 Falconiformes Accipitridae NT Declining
89
Natural Urban
Monticola saxatilis 15 0 Passeriformes Muscicapidae LC Depleted
Monticola solitarius 13 4 Passeriformes Muscicapidae LC Depleted
Montifringilla nivalis 9 0 Passeriformes Passeridae LC Secure
Morus bassanus 2 0 Pelecaniformes Sulidae LC Secure
Motacilla alba 67 42 Passeriformes Motacillidae LC Secure
Motacilla cinerea 55 25 Passeriformes Motacillidae LC Secure
Motacilla flava 47 28 Passeriformes Motacillidae LC Secure
Muscicapa striata 70 40 Passeriformes Muscicapidae LC Depleted
Neophron percnopterus 7 0 Falconiformes Accipitridae EN Endangered
Netta rufina 18 1 Anseriformes Anatidae LC Secure
Nucifraga caryocatactes 22 3 Passeriformes Corvidae LC Secure
Numenius arquata 25 4 Charadriiformes Scolopacidae NT Declining
Nycticorax nycticorax 15 4 Ciconiiformes Ardeidae LC Depleted
Oenanthe hispanica 7 0 Passeriformes Muscicapidae LC Depleted
Oenanthe oenanthe 42 22 Passeriformes Muscicapidae LC Declining
Oriolus oriolus 60 31 Passeriformes Oriolidae LC Secure
Otis tarda 4 0 Gruiformes Otididae VU Vulnerable
Otus scops 16 7 Strigiformes Strigidae LC Depleted
Oxyura leucocephala 2 0 Anseriformes Anatidae EN Vulnerable
Pandion haliaetus 12 0 Falconiformes Accipitridae LC Rare
Panurus biarmicus 19 4 Passeriformes Timaliidae LC Secure
90
Natural Urban
Parus ater 59 32 Passeriformes Paridae LC Stable
Parus caeruleus 66 43 Passeriformes Paridae LC Secure
Parus cristatus 50 27 Passeriformes Paridae LC Declining
Parus major 72 47 Passeriformes Paridae LC Increasing
Parus montanus 52 25 Passeriformes Paridae LC Secure
Parus palustris 56 27 Passeriformes Paridae LC Decreasing
Passer domesticus 58 44 Passeriformes Passeridae LC Declining
Passer montanus 59 36 Passeriformes Passeridae LC Declining
Pelecanus crispus 1 0 Pelecaniformes Pelecaniidae VU Rare
Perdix perdix 50 26 Galiiformes Phasianidae LC Vulnerable
Pernis apivorus 45 17 Falconiformes Accipitridae LC Secure
Petronia petronia 11 0 Passeriformes Passeridae LC Secure
Phalacrocorax carbo 16 4 Pelecaniformes Phalacrocoracidae LC Secure
Philomachus pugnax 6 4 Charadriiformes Scolopacidae LC Declining
Phoenicurus ochruros 54 36 Passeriformes Muscicapidae LC Secure
Phoenicurus phoenicurus 54 34 Passeriformes Muscicapidae LC Depleted
Phylloscopus bonelli 23 4 Passeriformes Sylviidae LC Declining
Phylloscopus collybita 70 36 Passeriformes Sylviidae LC Secure
Phylloscopus sibilatrix 54 29 Passeriformes Sylviidae LC Declining
Phylloscopus trochiloides 6 4 Passeriformes Sylviidae LC Secure
Phylloscopus trochilus 53 34 Passeriformes Sylviidae LC Secure
91
Natural Urban
Pica pica 59 45 Passeriformes Corvidae LC Secure
Picoides tridactylus 6 0 Piciformes Picidae LC Depleted
Picus canus 36 15 Piciformes Picidae LC Depleted
Picus viridis 58 36 Piciformes Picidae LC Stable
Platalea leucorodia 2 1 Ciconiiformes Threskiornithidae LC Rare
Plegadis falcinellus 3 0 Ciconiiformes Threskiornithidae LC Declining
Pluvialis apricaria 6 0 Charadriiformes Charadriidae LC Secure
Podiceps cristatus 46 22 Podicipediformes Podicipedidae LC Secure
Podiceps grisegena 16 4 Podicipediformes Podicipedidae LC Secure
Podiceps nigricollis 24 7 Podicipediformes Podicipedidae LC Secure
Porphyrio porphyrio 2 1 Gruiformes Rallidae LC Secure
Porzana parva 15 2 Gruiformes Rallidae LC Secure
Porzana porzana 25 9 Gruiformes Rallidae LC Secure
Prunella collaris 15 0 Passeriformes Prunellidae LC Secure
Prunella modularis 66 32 Passeriformes Prunellidae LC Secure
Pterocles orientalis 4 0 Columbiformes Pteroclididae LC Declining
Puffinus puffinus 2 0 Procellariiformes Procellariidae LC
Pyrrhocorax graculus 11 0 Passeriformes Corvidae LC Secure
Pyrrhocorax pyrrhocorax 11 1 Passeriformes Corvidae LC Declining
Pyrrhula pyrrhula 54 27 Passeriformes Fringillidae LC Secure
Rallus aquaticus 46 19 Gruiformes Rallidae LC Secure
92
Natural Urban
Recurvirostra avosetta 10 3 Charadriiformes Recurvirostridae LC Secure
Regulus ignicapilla 50 25 Passeriformes Reguliidae LC Stable
Regulus regulus 57 34 Passeriformes Reguliidae LC Decreasing
Remiz pendulinus 38 21 Passeriformes Remizidae LC Increasing
Riparia riparia 41 22 Passeriformes Hirundinidae LC Depleted
Rissa tridactyla 4 0 Charadriiformes Laridae LC Secure
Saxicola rubetra 54 18 Passeriformes Muscicapidae LC Secure
Saxicola torquatus 56 21 Passeriformes Muscicapidae LC Secure
Scolopax rusticola 37 13 Charadriiformes Scolopacidae LC Declining
Serinus serinus 55 36 Passeriformes Fringillidae LC Secure
Sitta europaea 64 33 Passeriformes Sittidae LC Secure
Somateria mollissima 4 0 Anseriformes Anatidae LC Secure
Stercorarius parasiticus 1 0 Charadriiformes Stercorariidae LC Secure
Sterna albifrons 12 5 Charadriiformes Laridae LC Secure
Sterna hirundo 26 10 Charadriiformes Laridae LC Secure
Streptopelia decaocto 52 41 Columbiformes Columbidae LC Secure
Streptopelia turtur 62 28 Columbiformes Columbidae LC Declining
Strix aluco 59 28 Strigiformes Strigidae LC Secure
Sturnus unicolor 9 1 Passeriformes Sturnidae LC Secure
Sturnus vulgaris 61 45 Passeriformes Sturnidae LC Declining
Sylvia atricapilla 75 43 Passeriformes Sylviidae LC Secure
93
Natural Urban
Sylvia borin 64 33 Passeriformes Sylviidae LC Secure
Sylvia cantillans 13 5 Passeriformes Sylviidae LC Secure
Sylvia communis 68 37 Passeriformes Sylviidae LC Secure
Sylvia conspicillata 8 0 Passeriformes Sylviidae LC
Sylvia curruca 51 33 Passeriformes Sylviidae LC Secure
Sylvia hortensis 9 0 Passeriformes Sylviidae LC Depleted
Sylvia melanocephala 15 8 Passeriformes Sylviidae LC Secure
Sylvia nisoria 23 12 Passeriformes Sylviidae LC Secure
Sylvia undata 15 1 Passeriformes Sylviidae NT Depleted
Tachybaptus ruficollis 55 23 Podicipediformes Podicipedidae LC Secure
Tachymarptis melba 14 1 Apodiformes Apodidae LC Secure
Tadorna tadorna 15 4 Anseriformes Anatidae LC Secure
Tetrao tetrix 22 1 Galiiformes Phasianidae LC Depleted
Tetrao urogallus 16 0 Galiiformes Phasianidae LC Secure
Tetrax tetrax 6 0 Gruiformes Otididae NT Vulnerable
Tichodroma muraria 9 1 Passeriformes Sittidae LC Secure
Tringa glareola 6 2 Charadriiformes Scolopacidae LC Depleted
Tringa ochropus 16 4 Charadriiformes Scolopacidae LC Secure
Tringa totanus 25 8 Charadriiformes Scolopacidae LC Declining
Troglodytes troglodytes 69 37 Passeriformes Troglodytidae LC Secure
Turdus iliacus 12 8 Passeriformes Turdidae LC Secure
94
Natural Urban
Turdus merula 75 45 Passeriformes Turdidae LC Secure
Turdus philomelos 68 37 Passeriformes Turdidae LC Secure
Turdus pilaris 45 28 Passeriformes Turdidae LC Secure
Turdus torquatus 21 1 Passeriformes Turdidae LC Secure
Turdus viscivorus 60 26 Passeriformes Turdidae LC Secure
Tyto alba 45 22 Strigiformes Tytonidae LC Declining
Upupa epops 40 16 Coraciiformes Upupidae LC Declining
Uria aalge 2 0 Charadriiformes Alcidae LC Secure
Vanellus vanellus 58 28 Charadriiformes Charadriidae LC Vulnerable
Xenus cinereus 0 2 Charadriiformes Scolopacidae LC Secure
95
Appendix E. Ultrametric tree of the molecular phylogeny estimated for the 157 Bavarian native breeding bird
species, according to the method used to obtained the Global Phylogeny of Birds by Jetz et al. (2012b). For
more details see Methods.
96
Appendix F. Ultrametric tree of the molecular phylogeny estimated for the 297 European-wide assemblage
native breeding bird species, according to the method used to obtained the Global Phylogeny of Birds by
Jetz et al. (2012b). For more details see Methods.
97
a
98
Variable Number Variable measured
Bio 1 Annual mean temperature (°C)
Bio 2 Mean diurnal range (mean of monthly Max temp – Min temp) (°C)
Bio 3 Isothermatity [(Bio 2 / Bio 7) *100]
Bio 4 Temperature seasonality (standard deviation*100)
Bio 5 Maximum temperature of warmest month (°C)
Bio 6 Minimum temperature of coldest month (°C)
Bio 7 Temperature annual range (Bio 5 – Bio 6) (°C)
Bio 8 Mean temperature of wettest quarter (°C)
Bio 9 Mean temperature of driest quarter (°C)
Bio 10 Mean temperature of warmest quarter (°C)
Bio 11 Mean temperature of coldest quarter (°C)
Bio 12 Annual precipitation (mm)
Bio 13 Precipitation of wettest month (mm)
Bio 14 Precipitation of driest month (mm)
Bio 15 Precipitation seasonality (Coefficient of Variation)
Bio 16 Precipitation of wettest quarter (mm)
Bio 17 Precipitation of driest quarter (mm)
Bio 18 Precipitation of warmest quarter (mm)
Bio 19 Precipitation of coldest quarter (mm)
Appendix H. Biplots of the principal component analysis of the 19 bioclimatic variables (See
Table above) within the WordClim databse across grids in Bavaria (a) and European-wide
studies (b). Numbers presented are after the numbers of grids/studies.
99

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UNIVERSIDADE DE LISBOA

DEPARTAMENTO DE BIOLOGIA ANIMAL

Phylogenetic diversity of bird assemblages along different land-use

Margarida Ladeira Felício Gonçalves

DEPARTAMENTO DE BIOLOGIA ANIMAL

Phylogenetic diversity of bird assemblages along different land-use

Margarida Ladeira Felício Gonçalves

Dissertação orientada por Professor Dr. Roland Brandl, Departamento de

Mestrado em Biologia da Conservação

Embora ausente, o teu amor é a minha força, iluminando o meu

Universität Marburg for accepting me as his Master student, accompanying me

and teaching me throughout my ERASMUS placement. For his patience, total

availability, tolerance and dedication put upon my work.

company and friendship.

Um agradecimento ao Prof. Dr. Carlos Fernandes, da FCUL, por me ter

aceitado como sua orientanda, pela disponibilidade e dedicação, sobretudo nos

últimos meses de trabalho.

À sua equipa, queria agradecer pessoalmente ao André Silva pela

Aos meus irmãos, sobrinhos, cunhados e tios-avós pela força, apoio e

A todos os meus amigos que sempre me apoiaram.

Land-use intensification leads to the transformation of natural habitats

lineages. Following an evolutionary conservative approach, it is thus expected a

decrease of phylogenetic diversity of assemblages with increasing amount of

The present study evaluates the phylogenetic diversity (mean

phylogenetic and mean nearest neighbour distance) of native breeding bird

assemblages, considering the impact of land-use at regional and continental

for comparative reasons only. Regionally, across Bavaria, it was used

presence/absence maps within 34 km2 grids and different land-use types; in a

European-wide study it was analysed which factors contribute to this

assemblage composition, in general and which of them differ between natural

and antrhopogenic sites.

Overall it was found that the occurrence of bird species is driven by

regional analysis although species richness increased with increasing

percentage of anthropogenic land-use, both measures of phylogenetic diversity

showed an overall decrease. For European-wide analysis, overall diversity was

(g) and percentage of insectivores.

In a world where urbanization is increasing at an exponential rate it is

important to analyse and recognize what processes and which factors

shaping bird assemblages, acting as strong environmental filters, with a

powerful evolutionary force.

Key-words: phylogenetic structure, habitat filtering, urbanization, Generalized

A diversidade filogenética foi primariamente aplicada ao estudo da

conservação, sendo que as espécies prioritárias eram seleccionadas com base

na sua taxonomia. Hoje-em-dia, a importância da aplicação desta medida de

biodiversidade extende-se também ao estudo das comunidades, promovendo o

conhecimento dos processos evolutivos que levam à composição da sua

estrutura filogenética, bem como da interacção existente entre as espécies

exclusão competitiva entre espécies que limita a sua coexistência a longo

prazo; e (2) filtragem ambiental de espécies que persistem numa comunidade

devido à sua tolerância a factores abióticos. O primeiro processo leva a uma

sobre-dispersão filogenética, que tende a limitar a coexistência de espécies

relacionadas entre si, conduzindo a uma diversidade filogenética superior à

esperada sob condições aleatórias de estrutura comunitária. Pelo contrário, o

segundo processo leva a um agrupamento filogenético, no qual espécies

relacionadas tendem a coexistir e a diversidade filogenética revela-se menor

A intensificação do uso-da-terra tem, nas últimas décadas, levado à

transformação do ambiente em habitats antropogénicos, nomeadamente em

terrenos agrícolas e de pastoreio, ou num sentido mais extremo, em áreas

urbanas. É do conhecimento geral que tais habitats altamente modificados

actuam como filtros ambientais, selecionando apenas algumas espécies com

determinadas características que lhes permitem sobreviver e que pertencem

assim, a apenas deteterminadas linhagens. Seguindo um contexto evolutivo

a coexistir, levando à diminuição da diversidade filogenética, mesmo quando