Interactions between non-native parrot species their recipient environments

Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Interactions between non-native parrot species and their recipient environments Dailos Hernández Brito PhD Thesis Seville, 2020 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda RECOMMENDED CITATION: Hernández-Brito, D. (2020). Interactions between non-native parrot species and their recipient environments. PhD Thesis. University of Seville, Seville, Spain Cover and cover back photos are authorship of Dailos Hernández-Brito, who remains his intellectual owner. Any form of reproduction, distribution, public communication or transformation of the same without prior authorization of the author is forbidden. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Interactions and t Interaccione s y los Supervisors: Dr. José Luis Tutor: Dr. Javier Ba Department of Co ns between non - native parrot spe d their recipient environments - s entre especies de psitácidos exó los nuevos medios que ocupan Thesis for the degree of Doctor University of Seville Seville, 2020 Dailos Hernández Brito uis Tella Escobedo Dra. Martina Balbontín Arenas Conservation Biology pecies xóticos na Carrete Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Que no puede ser sino salvaje y desmedido Jean-Claude Nicolas Forestier Arquitecto paisajista (Durante el diseño del Parque de María Luisa) A mis padres Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda CONTENT SUMMARY / RESÚMEN ...............................................................................................1 THESIS INTRODUCTION .............................................................................................7 Biological invasions: an issue for global biodiversity ..........................................9 Novel interactions between non-native species and recipient communities .......11 Parrots, unintentional characters in the scenario of biological invasion .............12 Study species: the most successful invaders and other non-native parrot species ....................................................................15 Objectives, hypotheses and thesis structure ........................................................18 References ...........................................................................................................21 SECTION 1: BIOTIC INTERACTIONS IN CAVITY NESTER COMMUNITIES.....32 Chapter I: Crowding in the city: losing and winning competitors of an invasive bird ...............................................................................................34 References ...........................................................................................................61 Chapter II: Alien rose-ringed parakeets (Psittacula krameri) attack black rats (Rattus rattus) sometimes resulting in death ......................................68 References ...........................................................................................................75 Chapter III: Nest-site competition and killing by invasive parakeets cause the decline of a threatened bat population ................................................79 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Resúmen 4 RESÚMEN Las invasiones biológicas están entre las causas más importantes de pérdida de biodiversidad en todo el mundo, aunque las interacciones entre las especies invasoras, la biota nativa y el medio receptor son en su mayoría desconocidas. Su identificación puede mostrar cómo ciertas especies invasoras pueden establecerse exitosamente en nuevos ambientes y qué impactos ecológicos potenciales amenazan la biota nativa. Sin embargo, tanto las interacciones como sus impactos potenciales generalmente se pasan por alto, especialmente en las primeras etapas de la invasión, porque éstos pueden surgir entre especies que interactúan inesperadamente, por lo que su detección depende de estudios a largo plazo. Los loros (Orden Psittaciformes) son uno de los grupos más afectados por el comercio internacional de vida silvestre debido a su valor como mascotas, por lo que el 16% de las especies de loros presentan poblaciones establecidas fuera de su área de distribución nativa. Entre los loros no nativos, las cotorras de Kramer (Psittacula krameri) y argentina (Myiopsitta monachus) son las más exitosas para establecerse e invadir en nuevos ambientes, aunque se han producido numerosas introducciones en más de cuarenta países de todo el mundo, sus interacciones e impactos bióticos son poco conocidos. En esta tesis, combinamos diferentes observaciones (conductuales, de distribución espacial, ambientales y poblacionales) para evaluar el papel de las diferentes interacciones bióticas y sus impactos resultantes que surgen entre diferentes especies de loros no nativos (principalmente cotorras de Kramer y argentina), ambiente y biota receptores. Nuestros resultados muestran que las especies de loros no nativos desarrollan una amplia gama de interacciones bióticas (por ejemplo, competencia interespecífica por sitios de anidación, facilitación de recursos limitantes y mutualismos animal-planta) que pueden amenazar o beneficiar a varias especies coexistentes, tanto nativas como no nativas. Por lo tanto, la aparición de Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Resúmen 5 comportamientos adaptativos (por ejemplo, asociación e innovación en la anidación) en loros no nativos permite superar las barreras provenientes del ambiente receptor, como la resistencia biótica y la escasez de recursos limitantes, u otros obstáculos durante sus primeras etapas de establecimiento (por ejemplo, efectos Allee). En general, el amplio análisis y estudio a largo plazo de las interacciones bióticas aquí realizado muestran su importancia como uno de los mecanismos clave para el éxito del proceso de establecimiento e invasión en especies de loros no nativos. Además, sus impactos ecológicos resultantes son un riesgo para las especies nativas, especialmente las especies amenazadas, por lo que la implementación de planes de manejo eficientes es urgente para conservar la biodiversidad. Palabras clave: Psittaciformes; invasiones biológicas; competencia interespecífica; resistencia biótica; comensalismo; innovación comportamental; facilitación; hibridación; Efecto Allee; mutualismo planta-animal; dispersión de semillas; zoocoria; impacto; erradicación; planes de manejo; viabilidad poblacional Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 7 THESIS INTRODUCTION Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Thesis Introduction 9 Biological invasions: an issue for global biodiversity Biodiversity is resultant of natural history processes that operate in different space-time scales and influence on key factors from genetic, through species and environmental to ecologic by which assemble the biological heterogeneity (Walker 1992). Likewise, biodiversity influences on ecosystem functions and provides good and services to humanity (Cardinale et al. 2012). However, the disruption of human activity has driven deep alterations on these processes and consequently endanger the local and global biodiversity (Vitousek et al. 1997; Cardinale et al. 2012). In this sense, the increasing biodiversity loss and extinction rates during the last century are unprecedented in previous geological ages, resembling thus massive extinctions from fossil record evidences (Dirzo et al. 2014; Ceballos et al. 2015; Wilson 2016). Such human impact acceleration on biodiversity compromises its understanding and detection of its causes and consequences, which in its turn spoil efforts to preserve global biodiversity (Dawson et al. 2011). Thus, different disruptive human actions are the most important drivers of biodiversity loss such as habitat destruction, overexploitation, climate change and biological invasions (Sala et al. 2000; Williams et al. 2008), as well synergetic interactions between these drivers may increase further their impacts (Mantyka-pringle et al. 2012). Translocation of animal and plants by humans have occurred during different human migrations worldwide. Pimentel (2001) estimates that about 400,000 species have been removed from their natural geographical areas to another by human agency during the past 10,000 years. Although there is a considerable increasing in translocation rates since 18th century due to technological advances that have assisted the long-distance transport (Meyerson and Mooney 2007; Hulme 2009). By deliberately or accidentally transporting, a large number of species have been introduced in novel Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Thesis Introduction 10 environments, whereby these species have overcome geographical barriers and established non-native viable populations out of their native ranges (Blackburn et al. 2011; Seebens et al. 2017). The establishment and spread of these non-native species (i.e. biological invasion process) may involve impacts on native species, community dynamics and ecosystem functions (Clavero and García-Berthou 2005; Vilà et al. 2011), in addition to economic damages and health issues (Vitousek et al. 1996; Simberloff et al. 2013). Native species may be affected through negative impacts such as predation, competition, spread of diseases and hybridization (Mack et al. 2000; Money and Cleland 2001), hence, their consequences threat viability of native populations, even their extinction (Clavero et al . 2009; Bellard et al. 2016). Nevertheless, the role of factors that drive processes in which non-native species become invasive, establishing and spreading successfully in new geographical areas, it has been debatable among numerous hypotheses (Kolar and Lodge 2001; Catford et al. 2009; Blackburn et al. 2011). Overall, these key factors are mainly categorized in three groups that may show synergy between them and change their relevance regarding a specific stage of invasion process (i.e transport, introduction, establishment and spread) (Kolar and Lodge 2001; Duncan et al. 2003; Blackburn et al. 2011; Abellán et al. 2017). First, event-level factors regarding the number of introductions and individuals released across the invasion history (i.e propagule pressure) (Lockwood et al. 2005). Second, location-level factors attributed to recipient environment traits such as coexisting native species similar to non-native species and climate matching regarding their native ranges (Forsyth et al. 2004; Cardador et al. 2016). Third, species-level factors or advantageous traits that show non-native species such as high abundances range-wide of their natural distribution, fast demographic growth rates and behavioral flexibility (Sol et al. 2005; Blackburn et al. 2009; van Kleunen et al. 2010). Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Thesis Introduction 11 Whereas several biotic traits are considered in the understanding of biological invasion success, mechanisms of biotic interactions are relegated of impact studies on native communities, despite of their importance in shaping species distributions (Wisz et al. 2013). Both direct and indirect biotic interactions between non-native species and recipient communities are expected during different stages of invasion process, although it is widely unexplored their influence on success of establishment and spread of nonnative species (Catford et al. 2009), especially in vertebrates (Blackburn et al. 2011). Novel interactions between non-native species and recipient communities After their introduction in novel environments, non-native species may gain or lose biotic interactions in relation to interactions that are previously present in their natural distribution ranges (Catford et al. 2009). These interactions may influence on non-native and native populations, so they are also keys determinating the success or failure of establishment and invasion processes, both through the loss (e.g. Enemy Release and Missed Mutualisms Hypotheses; (Elton 1958; Alpert 2006)) and gain of interactions (e.g. Invasional Meltdown and Biotic Resistance Hypotheses; (Elton 1958; Simberloff and Von Holle 1999)). Moreover, non-native species are mostly generalist (Wolf et al. 1996; Cassey et al. 2004; Sol et al. 2005), hence, the emergence of novel interactions to replace generalist interactions may occur more rapidly than those which are restrictive such as specialist (Traveset and Richardson 2014). As examples of how novel interactions works both ways, on the one hand, recipient community may display biotic barriers for invasive species such as a high diversity of natural enemies (e.g. predators and competitors) that produce biotic resistance against invasion process (Vilà and Weiner 2004; de Rivera et al. 2005; Foelker et al. 2018). On the other hand, coexisting invasive species may develop beneficial interactions between them, facilitating thus their invasion process as well as their impacts on native community through invasional Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Thesis Introduction 12 meltdown processes (Adams et al. 2003; Johnson et al. 2009). All these interactions usually categorize regarding their effect on each interacting species, namely negative, positive and neutral interactions. (Simberloff and Von Holle 1999). Most studies about biotic interactions and associated ecological impacts on native species focus on negative facets such as predation, competition and parasitism (Blackburn et al. 2009; Clavero et al. 2009; Ruscoe et al. 2011). However, positive interactions, such as facilitation, may also develop between native and invasive species and benefit both species, although few studies have tested their population effects and invasion success (Richardson et al. 2000; Altieri 2010; Wright et al. 2016) and only recorded in plants and invertebrates. Biotic interactions play a role between different ecological hypotheses aimed at explaining biological invasions although a correct assessment of impacts on native communities may be challenging, chiefly overlooked impacts and long-term ecological effects (Strayer et al. 2006; Simberloff et al. 2013; Blackburn et al. 2014). Thus, it is essential the implementation of a holistic approach that displays the complexity of ecological interaction networks and that includes biotic interactions from invasive species and as well as life-history traits of invaders and recipient communities (Catford et al. 2009). Besides, the role of biotic interactions may hide due to other key factors if the focus is wrong depending on in which stage of invassion process is working (Abellán et al. 2017). Therefore, this holistic approach may be fundamental to predict impacts on native biota and environment, developing thus effective management plans against biological invasions. Parrots, unintentional characters in the scenario of biological invasion A wide variety of animal groups are removed from their native geographic ranges due to the international wildlife trade (Beissinger 2001; Oldfield 2003; Young et al. 2016), a Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Thesis Introduction 19 interactions showed by several non-native parrot species to infer how limitations from recipient environment may trigger behavioral innovations and strategies to avoid them, favoring thus the invasion process (Section 2). We focus on animal-plant mutualism in which non-native parrots species interact with recipient vegetal community and their potential effects on native flora (Section 3). Finally, we assess the feasibility and efficiency of a hypothetical management plan focused on an invasive parrot species (Section 4). This doctoral thesis is constituted by four sections that describe the framework, justification, specific hypotheses and predictions of main goals. Different analytical approaches have been used in each section, material and methods in this thesis are described in detail within each section. In the Section 1, we monitored an invasive population of rose-ringed parakeet in an urban park located in Seville, Spain, to test potential interspecific competition for nesting sites (tree cavities) between the cavity nester community. Besides, we censuded several monk parakeet populations in different invaded areas to assess its potential facilitation of nesting sites for coexisting cavity nester species. We address particular questions such as (i) whether rose-ringed parakeet exploits nesting resources unexploited by natives (opportunism hypothesis) or it may share same preferences to exploit resources outcompeting aggressively with the rest of cavity nesters (competition hypothesis) (Sol et al. 2012), (ii) how the distribution of rose-ringed parakeet nests and their interspecific aggressions influence on spatial nesting distribution of rest of cavity nesters, (iii) whether a coexisting threatened bat species, the greater noctule (Alcalde et al. 2016), is more sensitive to competition pressure or both species do not interaction as well as what impacts emerge on its population during a time window of 14 years regarding the growing rose-ringed parakeet population, (iv) whether monk parakeet also Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Thesis Introduction 20 facilitates alternative nesting sites with own built nests to several species in its invasive range, and (v) how tenant species and host monk parakeets benefit of resultant interspecific colonianism. Part of this work have been published in the next journals: PloS One (2014); Hystrix (2014), Royal Society Open Science (2018). In the Section 2, we recorded several adaptive strategies from several populations of three different non-native parrot species (monk parakeet, rose-ringed parakeet, and orange-winged amazon) to assess their role as facilitators of establishment and invasion processes of these parrot species against unsuitable environmental conditions. We address particular questions such as (i) whether the emergence of nesting association between monk parakeets and a native species, the white stork, depends of habitat features (rural and urban) and what benefits provide nesting association for both species, (ii) how predation pressure from native raptors determinates the success of establishment and spread of monk parakeet (Catford et al. 2009), (iii) whether the scarcity of available nesting sites is a barrier for spread of roseringed parakeet, (iv) how novel nesting behaviors from some individuals spread to the rest population and may assist the invasion process of rose-ringed parakeet (Lefebvre et al. 2004), (v) how an incipient non-native population of orange-winged amazon overcomes the mate shortage and other Allee effects (Courchamp et al. 1999), and (vi) whether hybridization between non-native parrot species may rescue populations in early stages of establishment (Luquet et al. 2011). Part of this work has been published in Frontiers in Zoology (2020). In the Section 3, we infer the role of non-native parrot species in animal-plant mutualisms, in which, parrots may play a role as seed dispersers by different mechanisms. We address particular questions such as (i) which seed dispersal mechanisms are widely present between different parrot species (i.e. epizoochory and Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Thesis Introduction 21 estomatochory) (Tella et al. 2015), (ii) whether parrots are promoters of secondary dispersal through food wasting when other animal species benefit of food facilitation, and (iii) how non-native parrot species may disrupt animal-plant interactions established between native biota and assist non-native plants in invasional meltdown processes (Traveset and Richardson 2014). Part of this work has been published in Scientific Reports (2019). In the Section 4, we assess through population viability analysis the efficiency of different management actions according to a hypothetical eradication plan focused on an invasive population of rose-ringed parakeet in Seville, Spain. Given ecological impacts showed in Section 1 and ethic discrepancies regarding management actions, it is urgently necessary a consensus based in scientific-technical terms. We address particular questions such as (i) what demographic impacts show different management actions (i.e. breeding failure or removing individuals) on rose-ringed parakeet population(ii) whether population size of invader determinates the effectiveness of alternative removing individuals methods (i.e. trapping and shooting), and (iii) whether eradication plans are more feasible than control plans as well as more effective at longterm approach both conservation and cost-benefit perspectives (Simberloff 2003). 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E., Genovesi, P., Hulme, P. E., Jeschke, J. M., ... & Bacher, S. (2017). No saturation in the accumulation of alien species worldwide. Nature communications, 8(1), 1-9. − Senar, J. C., Carrillo-Ortiz, J. G., Ortega-Segalerva, A., Dawson Pell, F. S. E., Pascual, J., Arroyo, L., ... & Hatchwell, B. J. (2019). The reproductive capacity of Monk Parakeets Myiopsitta monachus is higher in their invasive range. Bird Study, 66(1), 136-140. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Thesis Introduction 28 − Senar, J. C., Domènech, J., Arroyo, L., Torre, I., & Gordo, O. (2016). An evaluation of monk parakeet damage to crops in the metropolitan area of Barcelona. Animal Biodiversity and Conservation, 39(1), 141-145. − SIF (Seychelles Island Foundation) (2017) Ring-necked parakeet culled at Morne Blanc! SIF Newsletter 56:1–2 − Simberloff, D. (2003). How much information on population biology is needed to manage introduced species?. Conservation Biology, 17(1), 83-92. − Simberloff, D., & Von Holle, B. (1999). Positive interactions of nonindigenous species: invasional meltdown?. Biological invasions, 1(1), 21-32. − Simberloff, D., Martin, J. L., Genovesi, P., Maris, V., Wardle, D. A., Aronson, J., ... & Pyšek, P. (2013). Impacts of biological invasions: what's what and the way forward. Trends in ecology & evolution, 28(1), 58-66. − Sol, D., Bartomeus, I., & Griffin, A. S. (2012). The paradox of invasion in birds: competitive superiority or ecological opportunism?. Oecologia, 169(2), 553-564. − Sol, Daniel, et al. "Big brains, enhanced cognition, and response of birds to novel environments." Proceedings of the National Academy of Sciences 102.15 (2005): 5460-5465. − Stafford, T. (2003). Pest risk assessment for the monk parakeet in Oregon. Salem, Oregon, USA: Oregon Department of Agriculture. − Strayer, D. L., Eviner, V. T., Jeschke, J. M., & Pace, M. L. (2006). Understanding the long-term effects of species invasions. Trends in ecology & evolution, 21(11), 645-651. − Strubbe, D., & Matthysen, E. (2009a). Establishment success of invasive ring‐necked and monk parakeets in Europe. Journal of Biogeography, 36(12), 2264-2278. − Strubbe, D., & Matthysen, E. (2009b). Experimental evidence for nest-site competition between invasive ring-necked parakeets (Psittacula krameri) and native nuthatches (Sitta europaea). Biological Conservation, 142(8), 1588-1594. − Strubbe, D., Matthysen, E., & Graham, C. H. (2010). Assessing the potential impact of invasive ring‐necked parakeets Psittacula krameri on native nuthatches Sitta europeae in Belgium. Journal of Applied Ecology, 47(3), 549-557. − Tella, J. L. (2011). The unknown extent of ancient bird introductions. Ardeola, 58(2), 399-404. − Tella, J. L., & Hiraldo, F. (2014). Illegal and legal parrot trade shows a longterm, cross-cultural preference for the most attractive species increasing their risk of extinction. PLoS One, 9(9), e107546. − Tella, J. L., Baños-Villalba, A., Hernández-Brito, D., Rojas, A., Pacífico, E., Díaz-Luque, J. A., ... & Hiraldo, F. (2015). Parrots as overlooked seed dispersers. − Tella, J. L., F. Hiraldo, E. Pacífico, J. A. Díaz-Luque, F. V. Dénes, F. M. Fontoura, N. Guedes, and G. Blanco. 2020. Conserving the diversity of ecological interactions: the role of two threatened macaw species as legitimate dispersers of “megafaunal” fruits. Diversity 12:45. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 35 ABSTRACT Invasive species can take advantage of resources unexploited by natives (opportunism hypothesis) or they can exploit the same resources but more aggressively or efficiently (competition hypothesis), thus impacting native species. However, invasive species tend to exploit anthropogenic habitats that are inefficiently used by natives such as urban environments. Focusing on the ring-necked parakeet (Psittacula krameri), one of the most invasive birds worldwide, we combined observations of interspecific aggressions, species-specific cavity-nest preferences and the spatial distribution of the native cavitynesting vertebrate community to determine the invasion process as well as its potential impacts on native species in a Mediterranean city. Our results support the competition hypothesis, suggesting that ring-necked parakeets are outcompeting native species sharing nest-site preferences. Parakeets initiated and won most interspecific aggressions, which were directed towards competitors but also towards predators. This behaviour could explain the spatial arrangement of natives, with most bird species breeding close to parakeets possibly to take advantage of their effective antipredatory behaviour. However, temporal and spatial patterns of segregation suggest that a threatened bat species is negatively affected by parakeets. This demonstrates that common species gain benefits and threatened ones (in this study, a bat and possibly a falcon) lose nest sites due to invaders. Therefore, the conservation status of the native species that pay the costs of competition with invaders should be considered. This scenario of winners and losers may, however, shift towards more losers if the ringnecked parakeet population continues to grow, thus requiring close monitoring and control/eradication programs to avoid further impacts. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 36 INTRODUCTION Biological invasions are considered a major threat to global biodiversity, since invasive species may cause negative impacts on natives through increased predation risk, competition, hybridization or the spread of disease (Mack et al. 2000). At smaller scales, however, the relationship between invasive species and biodiversity measures is less clear (Rosenzweig 2001), as introduced species can contribute to species gain by its establishment (Sax et al. 2002), can reduce species richness through extinction processes (Claveri et al. 2009) or can have no detectable effects on native biota (Simberloff 1981). These different patterns may be explained by the nature of the invader (Sax and Gaines 2008) but also by the characteristics of the recipient community (Catford et al. 2009). In an opportunistic scenario (formally called the empty niche, the invasion window or the opportunity window hypotheses; Catford et al. 2009), invasive species are functionally different from species already present in the community and thus their entrance into a new environment can occur without the displacement or extinction of natives. Conversely, when exotic and native species exploit similar resources, the recipient community could resist an invasion as a result of competition that stems from high local diversity and low niche vacancy (MacArthur 1970; Hierro et al. 2005). However, if exotic species are able to out-compete natives by exploiting resources more efficiently or through aggressive behaviours, they can successfully invade the new area causing the displacement of the native competitor (van Wilgenburg et al. 2010). Urban environments represent a challenge to biodiversity, as not all native species inhabiting the surrounding rural habitats are able to colonize these areas (Carrete and Tella 2011). Different studies have found a reduction in richness and diversity of native species along urban gradients, often in parallel with increments in exotic invasive Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 37 ones (McKinney 2006). Thus, as cities expand across the globe, biological homogenization increases as a consequence of the widespread increment of urbanadaptable, often invasive species at the expense of native, often endemic ones (McKinney and Lockwood 1999). This pattern suggests that many exotic and native species may not compete in nature (Blackburn et al. 2009), as the former tend to be particularly abundant in habitats that are inefficiently used by natives, such as in urban environments (Sol et al. 2012). However, cities still serve as refuges and conservation areas for some endangered natives (Miller and Hobbs 2002), which might come into conflict with invasive species using highly similar resources. Ring-necked parakeets (Psittacula krameri) are native to Asia and Africa and have established non-native urban populations in at least 35 countries on five continents (Butler 2003). Although it is considered amongst the 100 worst alien species in Europe (http://www.europe-aliens.org/speciesTheWorst.do), its impact on native species remains unclear. The ring-necked parakeet requires medium-size (4–8 cm entrance size, Strubbe and Matthysen 2009) natural cavities or those excavated in trees by other species for breeding. Given the usual shortage of tree cavities (Newton 1998), especially in urban environments where decaying tree limbs are periodically removed in the interests of public safety (Davis et al. 2013), parakeets could outcompete native cavitynesting species in aggressive interactions and thus spread at the cost of the numbers and/or distribution of natives (competition hypothesis). Alternatively, if the native community is poor in secondary cavity nesters and/or the resource is not limited, the establishment of this invasive species could be facilitated by a high availability of nesting sites (opportunism hypothesis). Previous work has shown that parakeets can outcompete only one of the coexisting native cavity-nesting bird species in a central- European city (Strubbe and Matthysen 2007, 2009), but larger-scale studies comparing Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 38 areas occupied and not occupied by this invader suggest little or no impact on populations of native birds (Strubbe et al. 2010; Newson et al. 2011). The outcome of invasive-native competition could be context-dependent, being influenced by the availability of resources and the composition of the native community (Catford et al. 2009). Thus, answering similar questions but using different systems can help to make generalizations about processes from local patterns. Here, we combined observations of interspecific aggressive interactions, species-specific cavity-nest preferences and the spatial distribution of cavities available and used by each species to infer the process as well as the consequences of ring-necked parakeet invasions in a Mediterranean city. Results show a complex scenario where, although ring-necked parakeets outcompete native species in aggressive encounters, most natives seem to benefit from the effective anti-predator behaviour of parakeets. Conversely, some threatened native species can be displaced by parakeets, resulting in a dynamic process of winners and losers linked to the population growth of the invader. MATERIAL AND METHODS Ethics Statement Field work conducted here was not invasive and did not require the manipulation of live animals. Therefore, this work did not require specific permits by the relevant Spanish authorities. Study System The ring-necked parakeet was a commonly traded wild species for the Spanish cagebird market (Carrete and Tella 2008a) and a number of urban populations arose largely from accidental escapes from cages (Martí and del Moral 2003). This study focuses on Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 39 the city of Seville (southern Spain), where the first records of the species date back to the early 1990’s and the initially small population sharply increased (Authors’ unpublished data), reaching ca. 1,000 individuals in 2011 (P. Edelaar com. pers.). We conducted the first breeding census of the species in the city of Seville from March to July 2013. We first located potential breeding areas where the species was present taking advantage of its conspicuous behaviour. Then, we monitored the available cavities to assess whether or not they were occupied by parakeets based on the observation of adults entering a minimum of 10 times on different days, the vocalizations of chicks inside the nest, and/or the observation of juveniles at the entrance. We located 216 active nests, 159 (73.6%) in an urban park (the María Luisa Park; 37° 22′ 31.57" N, 5° 59′ 19.59" W) and the rest forming smaller breeding nuclei in scattered groups of trees or, more rarely, in buildings throughout the city. María Luisa is the largest park located in the core of the city, comprising a 40 ha wooded area with a variety of tree species, most of them exotics such as Platanus sp., Eucalyptus sp. or Gleditsia triacanthos. The park is completely surrounded by streets with moderate to high traffic intensity. Availability and Occupancy of Tree Cavities The assessment of the availability of tree cavities and their occupancy by parakeets and native species was restricted to María Luisa Park to avoid potential biases when analysing interspecific competition (e.g., small groups of trees outside of the park occupied by parakeets could not be occupied by some native species because they did not offer sufficient foraging habitat). We GPS located (±3 m) all tree cavities that we were able to visually inspect in trees located within the park by using 10×50 binoculars. In each case, we identified the tree species and estimated the height of the cavity above ground (in m) and the width of its entrance (in cm). The entrances of cavities were Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 40 categorized as small, medium or large (<4 cm, 4–8 cm, and >8 cm, respectively) according to previous studies, which showed the preference of parakeets for cavities with entrances between 4 and 8 cm width (Strubbe and Matthysen 2007; Czajka et al. 2011). Cavities located at <2.5 m above ground were not considered for analyses since their accessibility to humans would preclude its use (none were occupied by native or exotic species), thus biasing results. From January to August (covering the entire breeding season of native and exotic species), we repeatedly visited and observed at a distance (for a minimum of 10 min) each cavity on at least 10 different days during daylight hours to assess whether or not it was occupied and by what species, devoting 48 days (202.5 hours) of field work. The close proximity of many trees with cavities often allowed us to monitor several trees simultaneously. A cavity was considered as occupied by a given bird species when we observed adults entering a minimum of 10 times on different days, heard chicks inside, or observed juveniles at the entrance. In addition, María Luisa Park is also inhabited by the greater noctule (Nyctalus lasiopterus). This cavity-breeding forest species is the largest European bat (averaging 48 g; Ibáñez et al. 2001), and the whole population living in and around Seville (roughly estimated at ca. 500 individuals in 2003–2004) gathers to breed and roost communally in the tree cavities of this park (Popa-Lisseanu et al. 2008; Ibáñez et al. 2009). To identify the cavities used by greater noctules, we detected their presence using an ultrasound detector (Pettersson D 230) and observing bats leaving tree cavities at sunset. Greater noctules, like other forest bats, form fission-fusion societies that switch roosts every few days, so each bat colony can control a large number of roosts of which only a few are occupied at a specific time (Popa-Lisseanu et al. 2008). Thus, using previous information on radio-tracked individuals (Popa-Lisseanu et al. 2008), we considered that a tree was not used by greater noctules during spring-summer 2013 if we did not Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 41 observe activity during any of our 10 spaced visits. Using information on trees used by noctules during 2003–2004, we also tested for changes in their use in relation to the current nesting spatial distribution of ring-necked parakeets. These trees were located after monitoring 27 noctules through radio-tracking to study the spatial pattern of tree use by the species, finding that cavities located in 75 trees were alternatively used as roost sites ten years ago (Popa-Lisseanu et al. 2008). It is worth noting that the different methodologies used to identify occupied trees could produce false cases of inoccupation in both 2003–2004 (a larger period of time monitoring 27 individuals) and 2013 (a shorter period of time monitoring all tree cavities). However, it may also just produce statistical noise making our estimates conservative. Spatial Distribution of Occupied Cavities The occupancy of a particular tree cavity by a given species could be influenced by the spatial distribution of cavities occupied by the same and/or other species, driven not only by competition but also by conspecific and heterospecific attraction processes. We thus obtained the distance from each occupied cavity to the nearest cavity occupied by conspecifics and heterospecifics (nearest neighbour distance) as well as the corresponding nest aggregation indexes. These aggregation indexes were obtained as the relative position of each occupied cavity within the whole distribution of all cavities occupied by conspecifics or heterospecifics in the park using ∑ exp (-dij), with (i≠j) where dij is the linear distance between occupied cavities i and j, j representing all occupied cavities (Moilanen and Hanski 1998). These variables were complementary measures depicting the social environment around each nest cavity at a landscape scale as well as the existence of close competitors. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 42 The spatial distribution of occupied cavities could also be influenced by habitat heterogeneity in the park. We considered the two main sources of habitat heterogeneity in our study area, i.e. the proximity to surrounding streets and forest cover. Noise from car traffic could alter song performance, reproductive success and even the spatial distribution of birds (Slabbekoorn and Peet 2003; Halfwerk et al. 2011). We therefore measured the linear distance from each cavity to the closest street using GIS tools (see below). On the other hand, species could differ in their preferences for forest coverage around cavities. We obtained forest cover by measuring it in a radius of 30 m around each GPS located cavity on a Google Map image taken in 2013 (Imagens ©2013 Cnes/Spot Image, DigitalGlobe, Instituto de Cartografía de Andalucía, map data ©2013 Google, based on BCN IGN Spain), using OpenLayers Plugin (1.1.0) applications in Q- GIS 1.8.0 (2008 Free Software Foundation, Inc). Forest cover was then scored into four main categories, namely 0–25%, 25–50%, 50–75%, and 75–100%. Interspecific Interactions We assessed interspecific interactions by randomly sampling the behaviour of different nesting ring-necked parakeets during a 15-minute period. We conservatively recorded the bird species present within a radius of 15 m around the focal parakeet, whether or not there was an aggressive interaction, what species started the attack, and which was the winner. To increase sample size without resampling the same individuals or the number of potential interacting species, observations were conducted in María Luisa Park as well as in other urban areas of Seville occupied by the species (see above), totaling 88 days (351.5 hours) of field work. These areas included the main parks of the city as well as a church (Divino Salvador) where ring-necked parakeets occupied cavities in walls for breeding, potentially competing there with lesser kestrels (Falco naumanni), a colonial falcon that usually breeds in urban buildings (Tella et al. 1996). Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 43 Statistical Analysis We employed Generalized Linear Models (GLM) implemented through the GENMOD procedure in SAS 9.2 (SAS Institute 2008) to ascertain which variables determined hole occupation, using the binomial error distribution (cavity occupied or not occupied by a given species) and the logistic link function. In a first set of models, we aimed to determine whether occupied and vacant cavities differed in their structural characteristics. Thus, we modelled the probability of occupancy as a function of the height of the cavity above ground (in its linear and quadratic forms), the entrance size, and the tree species. The resulting species-specific patterns of cavity preferences (see results) made it difficult to identify similarities (and thus opportunities for competition) between species. Therefore, we performed a categorical principal component analysis (CATPCA) on entrance size (since it is a categorical variable) and height above ground of the cavities occupied and took the scores of the obtained first dimension as a single compiling descriptor of the cavities used by each species. An ANOVA on these scores allowed us to identify differences in cavity preferences among species, and post-hoc Scheffe tests permitted us to establish homogeneous groups (i.e., species not differing in their preferences for particular cavity traits). In a second set of models, we assessed the spatial arrangement of each species regarding cavity traits, the distribution of both conspecifics and heterospecifics, and main habitat features (distance to the nearest street and forest cover) around each occupied and available (i.e., unoccupied) cavity, also using GLMs with a binomial error distribution and logistic link function. Exact binomial tests were used to assess whether the proportion of interspecific encounters ending in aggressions, the proportion of aggressions initiated by ring-necked parakeets, and the proportion of aggressions won by this species differed significantly from parity. To obtain interspecific patterns in the frequency of aggressions and their Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 44 outputs, we also used GLMs with a binomial error distribution and a logistic link function, fitting as explanatory variables the average body mass of the species interacting with ring-necked parakeets (obtained from Dunning 1993), their overlap in nest-site preferences (as a factor with levels ranging from 0– the interacting species was not a cavity-nester- to 3– maximum overlap in nest-site traits), and whether the interacting species was a potential predator of eggs, nestlings or adults. We expected that ring-necked parakeets would be more prone to attack those species with overlapping nest-site preferences and potential predators, and less prone to attack largerbodied species. A backward procedure was performed for GLM modelling, removing from full models those variables that were non-significantly associated with the response variable (p>0.05) to obtain minimum adequate models (MAM) (Sol et al. 2012). The resulting models did not show data overdispersion. We calculated the percentage of deviance explained as a measure of the variance explained by each MAM. RESULTS Occupancy of Tree Cavities We recorded 1,086 cavities in 435 trees located within María Luisa Park during the 2013 breeding season. Cavities were located at an average height above ground of 13.09 (SD 5.51) m, and the commonest cavity entrances (47%) were of intermediate size (4–8 cm). Most cavities (62.2%) were located in London plane trees (Platanus × acerifolia) probably because it is the most abundant species within the park but is also the species with highest number of available cavities. A total of 10 species were found occupying 525 cavities (Table 1), including 9 bird and one bat species. Two bird species were exotics, i.e. the ring-necked parakeet Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 51 encounters (Table 5), the probability that an encounter ended in aggression increased with the interspecific overlap in nest type preferences and the body mass of the interacting species, and decreased with the number of individuals of the interacting species involved. The probability that an aggression was initiated by ring-necked parakeets decreased with the body mass of the interacting species. Finally, the probability that a fight was won by ring-necked parakeets was greater if they initiated the attack but decreased when the interacting species was a potential avian predator. Nonetheless, ring-necked parakeets won 25–100% of the aggressions directed towards different predator species (Figure 2). Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 52 Figure 2. Percentage of encounters with ring-necked parakeets that ended in aggressions (white bars), and percentage of aggressions initiated (grey bars) and won by ring-necked parakeets (black bars). The number of recorded encounters is shown in brackets. Table 5. GLMs explaining the probability that an interspecific encounter ended in aggression (Aggression), whether the aggression was initiated by ring-necked parakeets (Fight initiation) and was won by ring-necked parakeets (Win fight). Nesting overlap N Body mass Attack initiation Predator % deviance Aggression 78.83 (+)*** 17.13 (-)*** 27.24 (-)*** 19.32 Fight initiation 21.89 (-)*** 9.58 Win fight 24.7 (+)*** 27.89 (-)*** 37.02 The retained explanatory variables were the interspecific overlap in nest types (Nesting overlap), the number of individuals of the interacting species present in the encounters (N), the average body mass of the interacting species (Body mass), whether or not ring-necked parakeets initiated the aggression (Attack initiation), and whether or not the interacting species is a potential predator of birds. χ 2 values are given for each variable. Signs between brackets indicate positive or negative effects of the explanatory variables. ***: p<0.001; % dev: percentage of deviance explained. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 53 DISCUSSION Opportunism or Competition? The successful establishment of exotic species in novel habitats constitutes a poorly understood paradox (Sax and Brown 2000). Recently, Sol et al. (2012) examined the invasion paradox by studying the use of food resources by invasive and native bird species in an Australian city, concluding that the success of invaders is explained by their capacity to exploit ecological opportunities that most native species rarely use. However, as the authors pointed out, competition over other resources, notably nesting sites, must be considered in further studies (Sol et al. 2012). We investigated two key aspects behind the establishment success of ringnecked parakeets on a relatively newly invaded urban area, namely: the way they shared nest-site resources with the recipient community and the aggressive interactions they experienced with other species. This approach allowed us to show that this species may invade new areas even when resources are not overabundant, thus not supporting the hypotheses proposing that saturated communities can halt biological invasions through competitive processes but rather supporting the competition hypothesis (instead of the opportunism hypothesis) for successful invasions (Catford et al. 2009). Although the availability of tree cavities was relatively high in the study area compared to other cities (Strubbe and Matthysen 2009; Orchan et al. 2013), the large populations sizes of different cavity-nester species together with the fact that the characteristics of unoccupied cavities differed from those of occupied ones suggest a shortage of suitable breeding sites for the native cavity-nesting community, coincident with the general pattern of competition found across cavity-nesting communities especially in urban environments (Newton 1994; Strubbe and Matthysen 2009; Orchan et al. 2013). Most of Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 54 the inability of the native community to resist the parakeet invasion may be due to the invader’s highly aggressive behaviour that allows it to out-compete natives, thus successfully occupying areas even when there is no superabundant or underexploited resources. Interestingly, we were able to separate species into three main functional groups based on species-specific nest site requirements, showing that parakeets fit into one of these groups. Thus, even when they may be interacting with many native species, they share important resources for population prospects (i.e., reproduction) with only some of them. However, ring-necked parakeets were aggressive (and won most aggressive encounters) not only towards those species sharing nest-site preferences (including two other exotic parrot species) but also towards others, even non cavitynesting species and avian predators. Therefore, the ring-necked parakeet has the potential to modify the numbers and spatial distribution of coexisting breeding species through behaviour-mediated competitive exclusion. Mechanisms behind the Spatial Arrangement of Species Habitat selection models, and species distribution models in their broader sense, are mathematical descriptions of biological patterns that are affected by environmental conditions and a multitude of direct and indirect interactions (Diamond 1975), thus inferring that causal links from observational data should be made with caution. Two species may co-occur if they share their habitat requirements, but also if they facilitate each other directly or indirectly. Conversely, species may appear to avoid each other if they show competitive exclusion but also if they have dissimilar habitat requirements. Although competition for cavities can trigger intraspecific negative interactions among individuals, we detected a general tendency among species to breed following a pattern of conspecific aggregation. This seems not to result from heterogeneities in habitat and nest site availability, aspects which were controlled for in statistical analyses, and thus Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 55 may rather be related to conspecific attraction processes as previously observed in many other colonial but also territorial species (e.g. Stamps 1988; Serrano et al. 2004). Different studies have shown that breeding in close proximity to conspecifics benefits breeders from earlier detection of predators, group defence, and dilution of predation (Brown and Brown 2001; Krause and Ruxton 2002; Serrano et al. 2005). Regarding the effects of the invasive species, we found that the spatial distribution of nesting ring-necked parakeets was important to explain the distribution patterns of all tree-cavity nester species of the recipient community while controlling for the main habitat features. However, the underlying putative mechanisms (attraction or segregation) were different among species. All bird species increased their likelihood of occupying cavities located close to parakeet nests and/or to high densities of parakeets. Positive co-occurrence patterns are indicative of heterospecific attraction (Sebastián- González et al. 2010), thus signaling the presence of direct or indirect species interactions. In our study system, a possible explanation for this association pattern could be found in the high aggressiveness of parakeets against avian predators. In fact, ring-necked parakeets may even communally attack predators, as we observed a flock of 60 parakeets mobbing a booted eagle (Aquila pennata) in María Luisa Park in 2008. Therefore, native species may choose breeding sites far enough from ring-necked parakeets (>15 m) to avoid aggressions but close enough to be rewarded by their effective anti-predator response, resulting in an active breeding association, which benefits the associated species (Blanco and Tella 1997). Conversely, the mutual spatial segregation between parakeets and noctules, not explained by habitat features, could be indicative of direct competition since they share their preferences for the same kind of cavities. The nocturnal behaviour of bats precluded the systematic observation of encounters with ring-necked parakeets, which would have been restricted to instances Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 56 when parakeets would enter bat cavities and inspect for potential nest sites. Although greater noctules aerially hunt small passerines when migrating at night (Ibáñez et al. 2001), they are not able to kill birds inside their nests (J. Juste com. pers.) and even less so a much larger species such as the ring-necked parakeet whose body mass (116 g) is more than twice that of the noctule (50 g). Given that parakeets won most aggressions when encountering larger-bodied competitors such as feral pigeons and even powerful jackdaws (Figure 2), they would be expected to also win most aggressive interactions with this much smaller bat species. Although little is known about the effects of aggressive species like parakeets on mammals that shelter and reproduce in hollows like bats, several authors suggested that they can evict them (Gebhardt 1996; Start 1998) and there is concern that ring-necked parakeets could cause the loss of suitable cavities for the noctule bat (Nyctalus noctula) in The Netherlands (Haarsma and van der Graaf 2009). In our study area, a greater noctule was fortuitously observed being aggressively expulsed from its cavity by a ring-necked parakeet in María Luisa Park in 2005 (E. Revilla com. pers.), and it could be expected that the strong beak of parakeets could seriously injure noctules to the point of killing or impeding their flight by irreversibly damaging their sensible patagium (J. Juste com pers.). Moreover, there is a published observation of a similar body-sized exotic parakeet (Superb Parrot, Polytelis swainsonii) killing the much larger red squirrel (Sciurus vulgaris, 295 g; Palomo et al. 2007) in Italy (Mori et al. 2013) and evidence of similar cases that might had been caused by ringnecked parakeets in France (Japiot 2005; Clergeau and Vergnes 2009). During this study we observed 11 instances of ring-necked parakeets (involving up to 10 individuals) attacking and mobbing black rats (Rattus rattus 180 g; Palomo et al. 2007), forcing them from the proximity of their nests. Since both rats and squirrels are predators of bird nests, including those of parakeets (Mori et al. 2013), these Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 57 observations also reinforce the potential benefits to other bird species of breeding close to parakeets. Impact: Winners and Losers in a Contemporary Invasion Process It is difficult to fully ascertain the ecological impacts of invaders, given the variety of potential impacts to be assessed, their subtle but pervasive effects, and the long time gaps between the introduction of an exotic species and its achievement of invasiveness and detectable impacts (Strayer et al. 2006; Simberloff et al. 2013). Although there are well-recognized cases of negative impacts of bird invasions in island environments (Lever 2005; Freed and Cann 2009), their impact on mainland environments have been less studied and much debated (Kumschick and Nentwig 2010; Bauer and Woog 2011; Strubbe et al. 2011), to the point of suggesting that introduced bird species should be managed before their negative impacts are proven (Edelaar and Tella 2012). Although ring-necked parakeets have been shown to outcompete a small cavitynesting native bird species in Brussels (Strubbe and Matthysen 2009), there is little evidence of its impact on native communities when comparing areas occupied or unoccupied by this invader (Strubbe et al. 2010; Newson et al. 2011). Our different approach, by recording the output of inter-specific aggressions and the spatial distribution of species in a Mediterranean city, suggests, however, that ring-necked parakeets may trigger strong effects on the native recipient communities, with both positive and negative responses depending on the native species considered. The spatial segregation of greater noctules and ring-necked parakeets together with the spatial patterns of trees abandoned during the last decade by noctules, its apparent population decrease and the parallel increase in the ring-necked parakeet population (Authors unpubl. data) suggest an active displacement exerted by the Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 58 invasive species. This is a matter of concern for this bat species, which shows a scattered distribution throughout Europe and is classified as Vulnerable in Spain, with María Luisa Park supporting its largest known colony (Ibáñez et al. 2004). Previous radio-tracking studies showed that greater noctules forage over large extensions of natural habitats (up to 40 km from the urban park (Popa-Lisseanu et al. 2009), including Doñana National Park and surrounding marshlands, but they return daily for roosting to Maria Luisa Park and no alternative refuges are known for this population (Popa- Lisseanu et al. 2008; 2009). This large population of greater noctules is therefore highly sensitive to any reduction in the availability of tree cavities caused by ring-necked parakeets. Given the scarcity of mature forests with large numbers of adequate cavities for the species, the other –although smallercolony of greater noctules known in South Spain is also located in an urban park (in Jerez de la Frontera, 77 km far from Seville) (Ibáñez et al. 2001). Although the presence of ring-necked parakeets is still anecdotic in this city, its population expansion might also pose threats to this bat population in the near future. Further studies are needed to deep on the population ecology and trends of greater noctule populations, and of other bat species (Haarsma and van der Graaf 2009), related to their coexistence with invasive parakeets. Another cause of concern is the fact that parakeets began to use wall cavities in 2011, breeding in three buildings in 2013, one of them located in the core of the city where there is also a colony of lesser kestrels. This colonial falcon suffered a drastic decline in Europe due to land-use changes that did not revert until recent years thanks to widespread conservation actions, including the provisioning of nest cavities (Catry et al. 2013). Lesser kestrels breeding in Seville have to forage far from their breeding colony (Tella et al. 1998) but gain benefits by the reduced predation risk in the city (Tella et al. 1996). Although their breeding success was linked to the quality of wall cavities (Negro Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 59 and Hiraldo 1993), the species was not constrained by nest-site availability or competition with feral pigeons and jackdaws in recent decades (Forero et al. 1996). However, the newly established ring-necked parakeet fought more than expected with lesser kestrels and won more than half of the aggressive encounters (Figure 2), while occupying only six wall cavities within the lesser kestrel colony. If the parakeet population continues to grow exponentially, it may pose a serious problem for urban lesser kestrels as well. In contrast to noctules and lesser kestrels, which are forced to forage far from the city, the abundance of food resources for ring-necked parakeets in the urban parks could reduce the energy they expend, allowing an increase in their breeding success and population growth (Strubbe and Matthysen 2011), thus reinforcing their competitive superiority. Both winner and loser species may result from anthropogenic-driven expansions of species (Carrete et al. 2010), some invaders even favoring whole communities of natives (Tablado et al. 2010). Our results suggest that the presence of nesting ringnecked parakeets may benefit several non-threatened native bird species, which may incur breeding advantages by exploiting their effective anti-predatory behavior. However, this situation could change in the near future if the ring-necked parakeet population continues to grow. This is already the most abundant species breeding in the park and is the only one able to enlarge tree cavities up to reaching its preferred size (4– 8 cm; Strubbe and Matthysen 2009; Czajka et al. 2011; this study), as has been shown in other urban parks (Orchan et al. 2013). In fact, 7 out of the 28 small-sized cavities (entrance <4 cm) were enlarged and occupied by parakeets during this study. Therefore, nest sites may become limited even for species using small-sized cavities such as tits and house sparrows. The latter species is a widespread commensal whose European Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter I 60 populations are now decreasing, thus drawing attention to its long-term conservation status (Robinson et al. 2005; Shaw et al. 2008). Conservation Implications We have shown potentially serious impacts of an invasive bird targeting species that are not easily monitored or that are not expected to interact with them, such as a forest bat and a colonial falcon nesting in buildings, thus highlighting the difficulties in assessing the entire set of impacts posed by invaders (Simberloff et al. 2013). The potential impact of ring-necked parakeets (Strubbe and Matthysen 2009), as well as of other parakeet species (Davis et al. 2013) thriving in urban habitats, has been often discounted since urban bird communities are usually composed by few, generalist and nonthreatened species (McKinney and Lockwood 1999; Sol et al. 2012). However, our case study suggests that urban ring-necked parakeets may be negatively affecting two threatened species, with some common species probably also affected in the near future if the parakeet population continues to grow. Therefore, both the conservation status of the native species with which the invader interacts as well as the population trends of the invader should be considered. Moreover, the positive population trends of ringnecked parakeets in Spain (authors' unpubl. data) suggests, as for monk parakeets (Muñoz and Real 2006), that the species could spread and invade rural habitats, as is already the case in central Spain (authors' unpubl. data). In such cases, parakeets would interact with a wider community of non-urban species and new impacts could arise, thus requiring a close monitoring of inter-specific interactions. As recommended for other invasive organisms (Simberloff et al. 2013), management of avian invasions should be undertaken before populations spread and actions become costly and even unaffordable (Edelaar and Tella 2012). In this regard, Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter II 68 Chapter II Alien rose-ringed parakeets (Psittacula krameri) attack black rats (Rattus rattus) sometimes resulting in death Dailos Hernández-Brito, Álvaro Luna, Martina Carrete, José L. Tella Hystrix It. J. Mamm,. 2014, 25(2):121–123 https://doi.org/10.4404/hystrix-25.2-10992 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter II 69 ABSTRACT The rose-ring parakeet (Psittacula krameri) is one of the most successful invasive birds in its establishment worldwide. Studies addressing its potential impact on native biota mostly focus on birds and little is known about how these and other parakeet species interact with native mammals. Here, we report 21 aggressions of rose-ringed parakeets towards black rats (Rattus rattus) in urban parks in Seville (Southern Spain) and Tenerife (Canary Islands). Either solitary parakeets or, more often, groups of up to 18 attacked rats when they climbed trees close to parakeet nests. Most attacks ended when the rats descended to the ground. However, in two instances (9.5% of the aggressions) the attacks resulted in the death of the rats as a result of falling to the pavement. These observations add further complexity to a biological invasion, where introduced parakeets have negative impacts on a predator and thus, some native bird species may benefit from their antipredator behavior. More attention should be paid to the interactions between native mammals and the non-native parakeets introduced worldwide. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter II 70 MAIN TEXT Biological invasions are considered among the main causes of biodiversity loss (Clavero and García-Berthou, 2005). Since the mid twentieth century, the introduction of exotic species has accelerated at an alarming rate (Hulme, 2009), mainly as a consequence of international trade (Meyerson and Mooney, 2007). Wild birds are among the most commonly traded vertebrate taxa (Carrete and Tella, 2008a), with a number of species having been accidentally introduced in recent times (Carrete and Tella, 2008b; Blackburn et al., 2010). Among these, parrots are one of the most heavily traded groups, mainly because of their attractiveness as pets (Tella and Hiraldo, 2014). Thirty-eight percent of birds offered for sale in Spanish pet shops between 2004 and 2005 were parrots belonging to 72 different species (Carrete and Tella, 2008b). Not surprisingly, several parrot species have established non-native populations worldwide (Lever, 2005; Menchetti and Mori, 2014). Introduced parrots may have a variety of ecological and economic impacts (Menchetti and Mori, 2014). Regarding ecological impacts, most research focuses on how introduced parrots affect native avifauna (e.g., Strubbe and Matthysen, 2007, 2009; Hernández-Brito et al., 2014). However, very little information is available on how they interact with native mammals. Mori et al. (2013) observed red squirrels preying on chicks at two rose-ringed parakeet (Psittacula krameri) nests, while an adult Barraband’s parakeet (Polytelis swainsonii) was responsible for the death of an adult red squirrel (Sciurus vulgaris) in Italy. Another report documents rose-ringed parakeets killing an adult red squirrel in France (Clergeau et al., 2009). More recently, Menchetti et al. (2014) recorded a fatal attack by a rose-ringed para keet of a Leisler’s bat (Nyctalus leisleri) in Italy. Here, we report several cases of rose-ringed parakeets attacking and even causing the death of black rats (Rattus rattus) in Spain. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter II 71 Observations of aggressions by non-native rose-ringed parakeets towards black rats were made in 2011, 2012, 2013 and 2014 in urban parks of Seville (Southern Spain), and Santa Cruz de Tenerife (Canary Islands). The two parks in Seville in which aggressions were recorded were Royal Alcázar (37°23'1.8''N / 5°59'29.6''W) and María Luisa Park (37°22'31.57''N / 5°59'19.59''W). Rose-ringed parakeets breed in both parks, with María Luisa Park holding the largest breeding nucleus in Seville (159 active nests in 2013; Hernández-Brito et al., 2014). Observations in Santa Cruz de Tenerife were made at García Sanabria Park (28°28'19''N / 16°15'13''W), where the breeding population of rose-ringed parakeets reached 12 pairs in 2014 (D. Hernández-Brito, unpubl. data). We observed 21 aggressions and attacks of rose-ringed parakeets on black rats, 16 in Seville and 5 in Tenerife. Interactions occurred when rats climbed trees where rose-ringed parakeets were nesting or trees in the vicinity of nests. When a rat was detected (Fig.1), rose-ringed parakeets first emitted loud alarm calls and then chased and attacked the rat using physical aggressions, sometimes biting it and producing visible wounds. All observed attacks were of solitary rats. Attacks were carried out by solitary parakeets or, more frequently, by groups of up to 18 individuals recruited from the surrounding breeding pairs (Fig.2A). In most cases, parakeets stopped the attack when rats escaped and descended or fell to the ground. Attacks caused the death of two rats in María Luisa Park. In the first event, a rat fell to the ground from a height of 15 meters when it was attacked by five rose-ringed parakeets in the treetops of a London plane. The rat fell directly onto the pavement, dying upon impact. The second rat died under similar circumstances; the rat was on a London plane with a rose-ringed parakeet nest and was attacked by six parakeets, failed to hold on to the branches and fell to the pavement from a height of 12 meters. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter II 72 Figure 1 – Instance of a black rat persecuted and attacked by a rose-ringed parakeet in Maria Luisa Park, Seville, May 25 2013. Darkness prevented a better-quality image (Photo: Dailos Hernández-Brito). The number of observations as well as the time of the day at which they were recorded suggests that attacks by rose-ringed parakeets towards rats are not anecdotal, but rather a difficult to observe behavior. Our observations were mostly carried out during scheduled field-work aimed to assess interactions between rose-ringed parakeets and native avifauna (Hernández-Brito et al., 2014), so most attacks were recorded in the morning or early afternoon (Fig.2B). In the twilight hours, rose-ringed parakeets gather in large flocks and leave for the roost (Pithon and Dytham, 1999; Hernández-Brito and Luna, unpubl. observ.). Only breeder females stay in their nest over night when their pairs return to the communal roost in the breeding season (Butler, 2003). Therefore, we expect more attacks would have been recorded if our observations included the crepuscular period before parakeets return to the roost, given the nocturnal behavior of rats (Hooker and Innes, 1995; Cox et al., 2000). Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter II 73 Figure 2 – Percentage of cases showing A) the number of rose-ringed parakeets involved in attacks on rats; and B) the time of day when attacks were recorded. Rats (Rattus sp.) are often invasive predators, and their negative effects on native and endemic avifauna in oceanic islands are well known Atkinson (1985); Traveset et al. (2009). On the other hand, urbanization changes the abundance and Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter II 74 richness of predator communities Chace andWalsh (2006), and thus rats can reach high population densities in the cities McKinney (2008). They can wipe out small populations of exotic parrots, as Scortecci (1953) recorded in the vicinity of a zoo in Italy, where a colony of monk parakeets (Myiopsitta monachus), the first established in this country (Spanò and Truffi, 1986), ceased to exist as a result of rat predation on parakeet eggs and chicks. At this point, and although we do not have direct evidence in the wild, black rats are potential predators of rose-ringed parakeet nests and even of adults (while incubating or sleeping, as has been recorded in captivity; J.L. Tella and M. Carrete, unpubl. observ.), so these aggressions may serve the same purpose as attacks on red squirrels, also known predators of parakeet nests (Mori et al., 2013). This aggressive behavior, along with the aggressiveness of rose-ringed parakeets towards avian predators, could explain why the nests of some bird species are aggregated in the proximity of nesting rose-ringed parakeets in Seville (Hernández-Brito et al., 2014). Although rose-ringed parakeets may displace (Hernández-Brito et al., 2014) and even kill coexisting bats, which compete for nesting cavities (Menchetti et al., 2014), other bird species may be favored by the antipredatory behavior of parakeets, thus increasing their breeding success. Three bird species (common blackbird Turdus merula, spotless starling Sturnus unicolor and collared dove Streptopelia decaocto) breeding in the proximity of rose-ringed parakeets joined the parakeets in chasing the rats in four cases. Together, these observations suggest that species benefit from other species’ aggressiveness toward predators (e.g., Blanco and Tella, 1997), and indicate that the impact of rose-ringed parakeets on native fauna may be both negative and positive depending on the species considered and its conservation status (Hernández-Brito et al., 2014). Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter II 75 Despite the fact that non-native populations of rose-ringed parakeets have spread throughout at least 35 countries (Butler, 2003), research aimed to assess their impact on native fauna has focused mostly on birds and has centered on only a few European cities (Strubbe and Matthysen, 2007, 2009; Orchan et al., 2013; Hernández-Brito et al., 2014). More attention should be devoted to their interactions with more elusive mammals, such as rats and bats, with a widespread occurrence in cities worldwide. ACKNOWLEDGEMENTS We thank the staff of García Sanabria Park, María Luisa Park and Royal Alcázar of Seville for allowing us access to these enclosures. Thanks to Emiliano Mori for comments that improved the manuscript. Field work was supported by Fundación Repsol and the Spanish Ministry of Economy and Competitiveness (project AIC-A- 2011-0706). The authors would like to acknowledge the support provided by the ParrotNet COST – European Cooperation in Science and Technology- Action, allowing fruitful discussions on data provided in this paper. REFERENCES − Atkinson, I. A. (1985). The spread of commensal species of Rattus to oceanic islands and their effects on island avifaunas. ICPB Tech Publ, 3, 35-81. − Blackburn, T. M., Gaston, K. J., & Parnell, M. (2010). Changes in non‐randomness in the expanding introduced avifauna of the world. Ecography, 33(1), 168-174. − Blanco, G., & Tella, J. L. (1997). Protective association and breeding advantages of choughs nesting in lesser kestrel colonies. Animal Behaviour, 54(2), 335-342. − Butler, C. J. (2003). Population biology of the introduced Rose-ringed Parakeet Psittacula krameri in the UK (Doctoral dissertation, University of Oxford). − Carrete, M., & Tella, J. L. (2008a). Non-native wildlife risk assessment: a call for scientific inquiry Response. − Carrete, M., & Tella, J. (2008b). Wild‐bird trade and exotic invasions: a new link of conservation concern?. Frontiers in Ecology and the Environment, 6(4), 207-211. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter II 76 − Chace, J. F., & Walsh, J. J. (2006). Urban effects on native avifauna: a review. Landscape and urban planning, 74(1), 46-69. − Clavero, M., & García-Berthou, E. (2005). Invasive species are a leading cause of animal extinctions. Trends in ecology & evolution, 20(3), 110. − Clergeau, P., Vergnes, A., & Delanoue, R. (2009). La perruche à collier Psittacula krameri introduite en Ile-de-France: distribution et régime alimentaire. Alauda (Dijon), 77(2), 121-132. − Cox M. P., Cox C. R. D., & Warren G. (2000). Use of habitat by the black rat (Rattus rattus) at North Head, New South Wales: an observational and experimental study. Austral Ecology, 25(4), 375-385. − Hernández-Brito, D., Carrete, M., Popa-Lisseanu, A. G., Ibáñez, C., & Tella, J. L. (2014). Crowding in the city: losing and winning competitors of an invasive bird. PloS one, 9(6), e100593. − Hooker, S., & Innes, J. (1995). Ranging behaviour of forest‐dwelling ship rats, Rattus rattus, and effects of poisoning with brodifacoum. New Zealand journal of zoology, 22(3), 291-304. − Hulme, P. E. (2009). Trade, transport and trouble: managing invasive species pathways in an era of globalization. Journal of applied ecology, 46(1), 10-18. − Lever, C. (2005). Naturalized Birds of the World. Poyser T. & Poyser, A.D, London. − McKinney M.L., 2008. Effects of urbanization on species richness: a review of plants and animals. Urban Ecosys. 11: 161-176. − McKinney, M. L. (2008). Effects of urbanization on species richness: a review of plants and animals. Urban ecosystems, 11(2), 161-176. − Menchetti M., Scalera R., Mori E., 2014. First record of a possibly overlooked impact by alien parrots on a bat (Nyctalus leisleri). Hystrix 25: 61-62. − Mori, E., Ancillotto, L., Menchetti, M., Romeo, C., & Ferrari, N. (2013). Italian red squirrels and introduced parakeets: victims or perpetrators?. − Meyerson, L. A., & Mooney, H. A. (2007). Invasive alien species in an era of globalization. Frontiers in Ecology and the Environment, 5(4), 199-208. − Orchan, Y., Chiron, F., Shwartz, A., & Kark, S. (2013). The complex interaction network among multiple invasive bird species in a cavity-nesting community. Biological Invasions, 15(2), 429-445. − Pithon, J. A., & Dytham, C. A. L. V. I. N. (1999). Census of the British Ringnecked Parakeet Psittacula krameri population by simultaneous counts of roosts. Bird Study, 46(1), 112-115. − Scortecci, G. (1953). Monaco (Myiopsitta monachus). In: Animali: come sono, dove vivono, come vivono. Milano, Italy: Labor Editors. − Spanò, S., & Truffi, G. (1986). Il Parrocchetto dal collare, Psittacula krameri, allo stato libero in Europa, con particolare riferimento alle presenze in Italia, e primi dati sul Pappagallo monaco, Myiopsitta monachus. Rivista italiana di Ornitologia, 56, 231-239. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 83 nesting species. There are no primary cavity-nesting species in this park (e.g. woodpeckers) that could excavate cavities, so all available cavities stem from tree decay. In addition, the land surrounding Seville is highly deforested and mostly devoted to agriculture, thus offering little habitat for forest-dwelling species. The greater noctule has a very scattered distribution throughout central and southern Europe, and Spain constitutes the main core area for the species (Alcalde et al. 2016). This species shows spatial sexual segregation (Ibáñez et al. 2009) and, during the breeding season, females gather in small maternity colonies (Alcalde et al. 2016). The María Luisa Park was thought to hold the largest local population of this species, as earlier studies conducted in just a sector of the park roughly estimated the presence of 500 adults (mostly females) (Ibáñez et al. 2009). Females form fission–fusion societies, resulting in several differentiated maternity colonies that occupy several cavities in different trees, with an average of 27 females simultaneously sharing a tree cavity (Popa-Lisseanu et al. 2008; Fortuna et al. 2009). Females may use different trees across the reproductive season and thus each maternity colony may occupy a minimum of 30 trees with adequate cavities, with the number of females using a particular cavity changing over time (Popa-Lisseanu et al. 2008). Nonetheless, females show long-term fidelity to the same groups of trees (some trees had been used for at least 14 years; Popa-Lisseanu et al. 2008). Females usually arrive at the park in March, giving birth to one or two pups in May–June. Although most of them leave the park from August to November, others stay year round (Popa-Lisseanu et al. 2008; Ibáñez et al. 2009). They have large foraging ranges, usually moving 15–40 km from the park in a single night to hunt insects and migrating songbirds (Popa-Lisseanu et al. 2009). Their annual survival rates were similar to those of other long-distance aerial-hawking bat species (Papadatou et al. 2012). Genetically, this population is closely related to the two closest colonies of Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 84 the species found in Jerez de la Frontera and Doñana National Park (Santos et al. 2016), located 60 km and 75 km from María Luisa Park, respectively. The rose-ringed parakeet is a successful invader in Europe. Their success is a result of the large numbers of individuals imported as cage birds, frequent accidental escapes or deliberated releases and niche similarity between areas of introduction and their native Asian habitats (Cardador et al. 2016; Abellán et al. 2017). The first introduction in Seville most probably occurred in 1992 with a very small group of individuals deliberately released in María Luisa Park. This is a highly sociable species that nests in cavities (mostly in trees but also in buildings; Hernández-Brito et al. 2014a) and produces one to four offspring per year. In María Luisa Park, most parakeets select tree cavities for nesting between late December and February and lay eggs mostly in March. There is, however, large variation in breeding phenology among pairs, with fledglings leaving the nests between May and early July. Moreover, newly formed immature pairs and groups of non-breeding individuals inspect and use tree cavities throughout the breeding season. Another eight cavity-nesting bird species are breeding in María Luisa Park (Hernández-Brito et al. 2014a). However, we have previously shown that the only species competing with noctules for tree cavities was the rose-ringed parakeet (Hernández-Brito et al. 2014a). Thus, the rest of the species were not considered in this study. Population monitoring Although our first studies on noctules in María Luisa Park date back to 1992 (Popa- Lisseanu et al. 2008), it was not until 2003 that we geolocated all trees used by the species in a single year. Unfortunately, we were not able to obtain a census of the whole Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 85 adult population of noctules. Obtaining accurate population sizes of forest bats forming fission–fusion societies, such as the greater noctule, is extremely difficult (Hayes et al. 2009; Kunz et al. 2009). In fact, there is no published information on population sizes for this or similar bat species. Therefore, we relied on temporal changes in the number of trees used by the species as a surrogate of changes in population size. We did not know the breeding population size of parakeets in 2003 as the species was still very scarce and little attention was paid to it because no impacts were expected at that time. We can, however, obtain a reliable estimate of the number of parakeet nests in 2003 from a demographic model built with the year of the first introduction, detailed population counts conducted since 2011 and breeding parameters obtained from the same population (Carrete et al. in prep.). In 2013 and 2016–2017, we simultaneously took a census of the number of nests of parakeets and the number of trees used by noctules in the park. Our monitoring programme extended from January to early August (covering the breeding season of both species), and observations were conducted in the morning (08.00–12.00 h) and afternoon (17.00–21.00 h). We GPS-located (±3 m) each year all tree cavities that we were able to visually inspect using 10 × 50 binoculars. We visually estimated the height of the cavity above ground (in m) and the width of its entrance (in cm), which was scored as small, medium or large (less than 4 cm, 4–8 cm and greater than 8 cm, respectively) (Hernández-Brito et al. 2014a). We did not consider the orientation of the cavities (Hernández-Brito et al. 2014a), as in other studies on nest-site competition between parakeets and native species (Orchan et al. 2013; Dodaro et al. 2014; Charter et al. 2016; Yosef et al. 2016; Mori et al. 2017), because orientation did not affect the selection of cavities in a previous study (Strubbe and Matthysen 2009). Our previous analyses showed a preference of both parakeets and noctules for cavities well above Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 86 ground (approx. 15 m) and with medium-sized entrances (Hernández-Brito et al. 2014a). Thereafter, we repeatedly visited and observed at a distance (for a minimum of 10 min) each cavity on at least 10 different days evenly spaced throughout the breeding season of the two species. Given that parakeets can enter cavities that are not used as nests (see above), we conservatively considered as active parakeet nests those cavities where we observed adults entering a minimum of 10 times on different days, heard chicks inside or observed juveniles at the entrance. Regarding the identification of trees used by noctules, we complemented the previous methodology for monitoring tree cavities with the use of an ultrasound bat detector (Pettersson D230) and systematic observations at sunset to observe noctules leaving tree cavities (Hernández-Brito et al. 2014a). Aggressive interactions Interactions between parakeets and noctules were observed during the monitoring activities described above. When we observed harassment and attacks of parakeets towards noctules present inside or in the entrance of their tree cavities, we recorded the duration of the aggression (in minutes), whether the noctule was expulsed and whether the parakeet later entered the cavity. Dead and injured noctules were also found, always under trees occupied by noctules. After unexpectedly encountering the first case of a dead noctule, we proceeded to record these events more systematically. We are convinced that many cases were overlooked because dead and injured noctules could be hidden by ground vegetation, scavenged by cats and rats or, as we later learned, recovered by people working in the park or visitors. The corpses we found were transported to the laboratory for a detailed visual examination of damage to the skin and bones. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 87 Spatial arrangement of noctules Each year (2013, 2016 and 2017) we geolocated the cavities used by parakeets and noctules as well as those that remained unoccupied. The occupancy of a particular tree cavity by noctules could be influenced by the availability of unoccupied cavities as well as by the spatial distribution of cavities occupied by the same and/or other species, driven not only by competition but also by conspecific attraction processes (Hernández- Brito et al. 2014a). We thus measured the Euclidean distance from each cavity to the nearest cavity occupied by noctules and parakeets (nearest-neighbour distance) as well as the corresponding aggregation indexes. Aggregation indexes were obtained as the relative position of each cavity within the whole distribution of all cavities occupied by conspecifics or heterospecifics using Σexp(−d ij ), with (i ≠ j), where d ij is the linear distance between cavities i and j, and j represents all occupied cavities (Moilanen and Hanski 1998). Nearest-neighbour distances and aggregation indexes are complementary and depict the social environment around each cavity at a landscape scale as well as the existence of close conspecifics and competitors in its proximity (Hernández-Brito et al. 2014a). The two main sources of habitat heterogeneity in our study area, i.e. the proximity to surrounding streets and forest cover, were not considered because our previous work showed no effects on the spatial arrangement of noctules and parakeets (Hernández-Brito et al. 2014a). We employed generalized linear models (GLMs) with a binomial error distribution and logistic-link function to ascertain factors explaining the probability of occupation of a cavity by noctules, fitting as explanatory variables its traits (entrance size and height above ground) and its nearest distance and aggregation to both noctules and parakeets. As values of nearest distance and aggregation for the same species were highly correlated (all p < 0.001), we alternatively included in models only one of these Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 88 spatial descriptors. Continuous variables were standardized for modelling. Models were separately built for the most distant years (2013 and 2017) from which we gathered spatial information from both species, using the Akaike information criterion corrected for small sample sizes (AICc) for model selection (Burnham and Anderson 2002). Within each set of candidate models, we calculated ∆AICci as the difference between the AICc of model i and that of the best-supported model (i.e. the model with the lowest AICc). Models within 2 AICc units of the best supporting model were considered as alternatives. We also quantified the plausibility of each model as being the best approximation using Akaike weights, w (Burnham and Anderson 2002), and performed model averaging (MuMIn package) to estimate the relative importance of all variables through the calculation of model-averaged estimates and confidence intervals (CIs) using the set of alternative models. A given effect received no, weak or strong support when the 95% CI for the coefficient estimate strongly overlapped zero, barely overlapped zero or did not overlap zero, respectively. We calculated the percentage of deviance explained by the best-supported models, obtained as 100 − (deviance of model i/deviance null model) * 100), for assessing their goodness of fit. All statistical analyses were conducted in R v. 3.1.2 (R Core Team 2013), and raw data used for analyses are provided in the electronic supplementary material. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 89 RESULTS Interspecific aggressions During the breeding seasons of 2016 and 2017, we recorded 36 aggressions of parakeets towards noctules in trees occupied by both species. Most aggressions (55.6%) were recorded in May and during the 3 h before sunset (83.3% of aggressions). All of these aggressions happened at the entrance of tree cavities, and in 16 cases, we observed parakeets attacking noctules with their beaks while emitting loud sounds (figure 1). These aggressions usually lasted from 1 to 25 min, reaching in one case up to 145 min (median: 13.13 min), and ended with the parakeets entering the tree cavity after the attacked noctule was expulsed and obliged to flee during the daytime. Once parakeets entered the tree cavity, we were not able to observe whether or not they attacked other noctules that may have been inside the cavity. In 20 other instances, parakeets ceased their aggression and left the tree at sunset without successfully expelling the noctules from their refuge cavities. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 90 Figure 1. Main picture: an adult female rose-ringed parakeet attacks a greater noctule at the entrance of a tree cavity used as a refuge by noctules in María Luisa Park, Seville, on 17 May 2016. One noctule was found dead under the same tree the next day, with wounds caused by parakeets. Accompanying pictures: details of different types of wounds caused by parakeets on noctules found dead under their tree refuges (photographs: Dailos Hernández-Brito). During the same period, we found 20 dead and two injured noctules under 18 different trees, all of them with active nests of parakeets. Seven of these noctules were found dead under the same tree where we recorded aggressions by parakeets the previous day. Dead noctules included three few-day-old pups, 10 lactating young and seven adults, as well as one pregnant female. Eight of these corpses were too putrefied Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 91 to allow us to observe wounds or other indications of attack (temperatures reached 33– 40°C during the study period, so corpses decomposed rapidly). The other 12 corpses were fresh enough to show clear wounds caused by the beaks of parakeets (figure 1). These wounds were present mainly on the wings, consisting of holes in the membranes (dactylopatagium and plagiopatagium areas), chafing on the skin covering the phalanges and forearm, breakages of phalanges, and wounds both on the upper side of the body and the abdomen. The two injured, but still living adult noctules showed similar wounds and were not able to fly or to climb the trunks of trees to return to their cavity refuges. We suspect that noctules were injured and killed when parakeets entered their cavities after expelling other noctules (see above), because pups and lactating young do not leave the cavities. We did not observe aggressions addressed to noctules by other species, neither did we find evidence of other species preying on or killing noctules in the park. Temporal trends in the occupation of trees Figure 2 shows the numbers of nests of parakeets and trees occupied by noctules in María Luisa Park from 2003 to 2017. Parakeets were scarce in this area in 2003 and, although we did not conduct a detailed census, our demographic-based estimation suggests that the local breeding population would have been as low as 13 nests at that time. Three annual detailed censuses conducted since 2013 showed an increase to 311 active nests in 2017 (figure 2). This means a 96% population increase from 2013 to 2017, and an estimated 2192% population increases from 2003 to 2017. Noctules showed an opposite trend: cavities used as refuges were found in 75 trees in 2003, the number decreasing to only 14 trees in 2017 (figure 2). This resulted in a 70% decrease between 2013 and 2017, and an 81% decrease between 2003 and 2017. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Figure 2. Changes in the n by greater noctules across y Spatial segregation between Figure 3 shows the spatial across years in the park, ind contraction of noctule refug nests. In 2013, the first y parakeet nests (n = 159 nes probability a tree cavity wa height above ground and to nests (table 1). In this yea medium- to largesize entra parakeet nest (table 2). In declined to 14 while the nu Sect e number of nests of rose - ringed parakeets and s years in María Luisa Park. een noctules and parakeets ial distribution of parakeet nests and noctule r indicating not only a reduction in the number b fuges along with an increase and spatial expan t year we recorded both noctule refuges (n nests) (figure 3), the best - supported model in was occupied by noctules was related to its en to the nearest distances to other noctule refug ear, noctules used cavities well above ground trances, close to other noctule refuges and fa r In 2017, when the number of trees occupied number of parakeet nests had increased to 31 ection 1: Chapter III 92 and trees occupied e refuges recorded r but also a spatial ansion in parakeet = 47 trees) and indicates that the entran ce size and fuges and parakeet nd level and with r from the nearest d by noctules had 311 (figure 3), the Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 99 predation rates is very limited (Barclay and Harder 2003). Moreover, the annual survival rates of noctules living in María Luisa Park before the population growth of parakeets was low (0.74) compared to other forest bat species that forage at shorter distances (Papadatou et al. 2012). Therefore, an increment in mortality rates caused by parakeets both on lactating and adult noctules would severely affect the population dynamics of this population and contribute to its dramatic decline. Unfortunately, we were unable to measure temporal changes in population size owing to the methodological difficulties of accurately estimating population sizes of forest bat species that form fission–fusion societies (Hayes et al. 2009; Kunz et al. 2009). However, it is reasonably expected that the population size would have declined in parallel with the 81% reduction in the number of tree cavities used. Population size, as well as social structure, is expected to be affected: females were originally distributed across the park forming differentiated, stable maternity colonies (Popa-Lisseanu et al. 2008; Fortuna et al. 2009; figure 3), something certainly disrupted given the scarcity and aggregation of the trees currently used (figure 3). This social disruption may have unexpected effects on the breeding biology of the species. Actions to reduce the population of parakeets and to provide artificial refuges for noctules are urgently needed. Without that, this population, which was once the largest known for this threatened bat species in its range (Fortuna et al. 2009), could be completely extinct within a few years. The provision of artificial refuges alone would be insufficient, given that noctules learn to use them slowly (C. Ibáñez 2003, unpublished data) and that the population size and direct impacts of parakeets are increasing at a much faster rate. An eradication plan of parakeets was planned for early 2017 by the city government of Seville, but this was cancelled owing to pressures by animal welfare associations, even though it was supported by a Spanish law which specifically deals Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 100 with the management of invasive species (Real Decreto 630/2013). Programmes for the control or eradication of invasive species often face public opposition (Blackburn et al. 2010), especially when dealing with charismatic species such as parakeets (Carrete and Tella 2008). Our work exemplifies the challenges in understanding the true ecological impacts of invasive birds. While the number of introduced non-native bird species increases worldwide (Dyer et al. 2017) and invasion risks increase in new regions (Cardador et al. 2017), the scientific community is only able to study a small fraction of these populations and their multifaceted potential impacts (Martin-Albarracin et al. 2015). Moreover, some impacts, such as those reported here, are unexpected and can be easily overlooked in the absence of long-term research. In this sense, we speculate whether other impacts on bats remain hidden. Many European bat species, some of them threatened, are closely linked to buildings and urban habitats (Ancillotto et al. 2015; Russo and Ancillotto 2015; Rydell et al. 2017)); meanwhile, parakeets are spreading across European cities (Cardador et al. 2016; Abellán et al. 2017). There is concern that parakeets could reduce the availability of suitable tree cavities for the noctule bat (Nyctalus noctula) in the Netherlands (Haarsma and van der Graaf 2013). One Leisler's bat (Nyctalus leisleri) was found killed by parakeets in Italy (Menchetti et al. 2014), and parakeets seem to be competing for greater noctule refuges in Jerez de la Frontera (I. Sánchez and D. Hernández-Brito 2017, personal observation). Parakeets can also compete for cavities in buildings that they use for nesting, which can be usurped from other bird (Hernández-Brito et al. 2014a) and bat species. One of the authors (D. Hernández-Brito) observed in Seville, on 7 July 2017, three parakeets attempting to force isabelline serotine bats (Eptesicus isabellinus) out from a wall cavity in a tall building during the daytime. Seventy-eight bats later flew out from that cavity at sunset. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 101 All of these observations can be considered as anecdotal and do not necessarily imply an impact on bat populations. Similarly, the first observation of parakeets harassing a noctule in María Luisa Park in 2005 (Hernández-Brito et al. 2014a) would not have indicated a population impact until the long-term research presented here. As a matter of concern, a recent meta-analysis shows that large impacts caused by invasive species can often be missed owing to small sample sizes, resulting in high Type II error rates and false certainty of no impact (Davidson and Hewitt 2014). Therefore, much more research is needed to properly assess the impact of parakeets and other invasive species on a variety of bat species (Welc and Leppanen 2017). ACKNOWLEDGEMENTS We thank the María Luisa Park staff for facilitating the work within the park. J. Nogueras helped with data management, and several people from the Department of Conservation Biology and Evolutionary Ecology of EBD-CSIC, students from Pablo de Olavide University and other volunteers helped to record field observations. E. Mori and an anonymous reviewer helped to improve this paper. This work was funded by the Organismo Autónomo de Parques Nacionales (MMA) of Spain (project 021/2002) to C.I., and Fundación Repsol, the Severo Ochoa Program (SVP-2014-068732) and Action COST ES1304 (ParrotNet) to D.H-B., J.L.T. and M.C SUPPLEMENTARY MATERIAL Appendix S1. (see Supplementary Material Section 1-Chapter III). Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 102 REFERENCES − Abellán, P., Tella, J. L., Carrete, M., Cardador, L., & Anadón, J. D. 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H., Betke, M., Hristov, N. I., & Vonhof, M. J. (2009). Methods for assessing colony size, population size, and relative abundance of bats. In Ecological and behavioral methods for the study of bats, 2nd edn (eds TH Kunz, S Parsons), pp 133–157. Baltimore, MD: The Johns Hopkins University Press. − Le Louarn, M., Couillens, B., Deschamps-Cottin, M., & Clergeau, P. (2016). Interference competition between an invasive parakeet and native bird species at feeding sites. Journal of ethology, 34(3), 291-298. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 104 − Martin-Albarracin, V. L., Amico, G. C., Simberloff, D., & Nuñez, M. A. (2015). Impact of non-native birds on native ecosystems: a global analysis. PLoS One, 10(11), e0143070. − Menchetti, M., & Mori, E. (2014). Worldwide impact of alien parrots (Aves Psittaciformes) on native biodiversity and environment: a review. Ethology Ecology & Evolution, 26(2-3), 172-194. − Menchetti, M., Mori, E., & Angelici, F. M. (2016). Effects of the recent world invasion by ring-necked parakeets Psittacula krameri. In Problematic wildlife (pp. 253-266). Springer, Cham. − Menchetti, M., Scalera, R., & Mori, E. (2014) First record of a possibly overlooked impact by alien parrots on a bat (Nyctalus leisleri). Hystrix 25, 61– 62. − Moilanen, A., & Hanski, I. (1998). Metapopulation dynamics: effects of habitat quality and landscape structure. Ecology, 79(7), 2503-2515. − Mori, E., Ancillotto, L., Menchetti, M., & Strubbe, D. (2017). ‘The early bird catches the nest’: possible competition between scops owls and ring‐necked parakeets. Animal Conservation, 20(5), 463-470. − Orchan, Y., Chiron, F., Shwartz, A., & Kark, S. (2013). The complex interaction network among multiple invasive bird species in a cavity-nesting community. Biological Invasions, 15(2), 429-445. − Papadatou, E., Pradel, R., Schaub, M., Dolch, D., Geiger, H., Ibañez, C., ... & Gimenez, O. (2012). Comparing survival among species with imperfect detection using multilevel analysis of mark—recapture data: a case study on bats. Ecography, 35(2), 153-161. − Pârâu, L. G., Strubbe, D., Mori, E., Menchetti, M., Ancillotto, L., Kleunen, A. V., ... & Clergeau, P. (2016). Rose-ringed Parakeet Psittacula krameri populations and numbers in Europe: a complete overview. The Open Ornithology Journal, 9(1). − Peck, H. L., Pringle, H. E., Marshall, H. H., Owens, I. P., & Lord, A. M. (2014). Experimental evidence of impacts of an invasive parakeet on foraging behavior of native birds. Behavioral Ecology, 25(3), 582-590. − Popa‐Lisseanu, A. G., Bontadina, F., & Ibáñez, C. (2009). Giant noctule bats face conflicting constraints between roosting and foraging in a fragmented and heterogeneous landscape. Journal of Zoology, 278(2), 126-133. − Popa-Lisseanu, A. G., Bontadina, F., Mora, O., & IbÁñez, C. (2008). Highly structured fission–fusion societies in an aerial-hawking, carnivorous bat. Animal behaviour, 75(2), 471-482. − R Core Team. 2013. R: a language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. See http://www.R- project.org/. − Russo, D., & Ancillotto, L. (2015). Sensitivity of bats to urbanization: a review. Mammalian Biology, 80(3), 205-212. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 1: Chapter III 105 − Rydell, J., Eklöf, J., & Sánchez-Navarro, S. (2017). Age of enlightenment: longterm effects of outdoor aesthetic lights on bats in churches. Royal Society open science, 4(8), 161077. − Santos, J. D., Meyer, C. F., Ibáñez, C., Popa‐Lisseanu, A. G., & Juste, J. (2016). Dispersal and group formation dynamics in a rare and endangered temperate forest bat (Nyctalus lasiopterus, Chiroptera: Vespertilionidae). Ecology and evolution, 6(22), 8193-8204. − Simberloff, D., Martin, J. L., Genovesi, P., Maris, V., Wardle, D. A., Aronson, J., ... & Pyšek, P. (2013). Impacts of biological invasions: what's what and the way forward. Trends in ecology & evolution, 28(1), 58-66. − Sol, D., González-Lagos, C., Lapiedra, O., & Díaz, M. (2017). Why are exotic birds so successful in urbanized environments?. In Ecology and conservation of birds in urban environments (pp. 75-89). Springer, Cham. − Strubbe, D., & Matthysen, E. (2009). Experimental evidence for nest-site competition between invasive ring-necked parakeets (Psittacula krameri) and native nuthatches (Sitta europaea). Biological Conservation, 142(8), 1588-1594. − Turbé, A., Strubbe, D., Mori, E., Carrete, M., Chiron, F., Clergeau, P., ... & Nentwig, W. (2017). Assessing the assessments: evaluation of four impact assessment protocols for invasive alien species. Diversity and Distributions, 23(3), 297-307. − Welch, J. N., & Leppanen, C. (2017). The threat of invasive species to bats: a review. Mammal Review, 47(4), 277-290. − Yosef, R., Zduniak, P., & Żmihorski, M. (2016, December). Invasive ringnecked parakeet negatively affects indigenous Eurasian hoopoe. In Annales Zoologici Fennici (Vol. 53, No. 5–6, pp. 281-287). Finnish Zoological and Botanical Publishing Board. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 124 SECTION 2: NOVEL INTERACTIONS AGAINST BIOTIC BARRIES Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 131 are also tolerated by white storks and use their nests as nesting substrates (Indykiewicz 2006). The large-bodied white stork has extremely low nest predation rates, as learned from long-term breeding monitoring programs in Spain (e.g.; Baos et al. 2012; Vergara et al. 2006), and predation of adults is anecdotal (Tobolka 2014). Therefore, smaller bird species may also associate with white storks to reduce predation risk (Bocheński 2005; Kosicki et al. 2007), as it has been demonstrated for other protective nesting associations (Quinn and Ueta 2008). Fig. 1 Parakeets and storks. Nests of monk parakeets (yellow arrows) associated with white stork nests. (Photos: D. Hernández-Brito) Here, we hypothesised that monk parakeets associate with storks not because of structural benefits but to obtain protection against predation, thus allowing their spread into rural habitats despite the existence of a large predator community. To test this hypothesis, we first compared the probability of parakeet-stork associations between rural and urban habitats, predicting that they should be more frequent in the former, where predators are more abundant than in the latter. We then assessed whether these associations occur at random, or if they can be explained by a combination of biotic and abiotic factors such as the proximity of the stork nest to another parakeet colony (conspecific density), the type of substrate (pylons, trees, or roofs), or the density of Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 132 predators in the surroundings (predation risk). We predicted that besides proximity to conspecifics, parakeets should breed preferentially in stork nests sited in pylons (thus reducing predation risk at the nest) in areas with a low density of aerial predators (thus reducing predation risk while foraging). The low density of predators in the surrounding of a nest ensures safe areas for parakeets to perform basic activities such as foraging. Nests in pylons, contrary to those located in trees or on roofs, are not accessible to mammalian or reptilian predators (Bocheński 2005), so the antipredatory effect of nesting with a stork is focused on aerial predators (i.e., raptors) and, thus, maximised. Finally, to discard the potential benefits derived from the nest structure per se, we evaluated the effect of nest abandonment by storks on the subsequent nest abandonment by parakeets. We predict that parakeets should abandon their nests after stork abandonment due to the disappearance of its protective effect against predators; otherwise, parakeets could associate with storks to simply take advantage of their nest structures. Complementarily, we compared the behaviour of parakeets toward approaching avian predators when breeding in association with storks or not. We predicted that parakeets should flush more frequently from raptors when breeding alone than when breeding with storks, where they can take advantage of the presence of the protective species, which can deter raptor attacks. MATERIAL AND METHODS Study area and fieldwork The study was carried out in an extensive area including the city of Madrid and its surrounding rural habitats along the Manzanares and Jarama rivers (Fig. (Fig.2),2), an area mostly devoted to irrigated, intensive agriculture (mainly maize and vegetables) and gravel extraction. In this area, raptors nest mostly in riparian forests, while white Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 133 storks nest in the same forest as well as on electric pylons and building roofs (Blanco 1994; 1996). During the breeding seasons (April–August) of 2014 and 2015, the study area was repeatedly visited to GPS-locate all nests of parakeets and storks present in the urban and rural habitats (Fig. (Fig.2),2), also recording the type of substrate in which the nests were located (tree, pylon or roof). Moreover, we monitored the community of medium-sized raptor species present in the study area, including the black kite Milvus migrans, the booted eagle Hieraaetus pennatus, the common buzzard Buteo buteo, the northern goshawk Accipiter gentilis, the red kite Milvus milvus and the peregrine falcon Falco peregrinus. Predation of monk parakeets by raptors has been recorded in its invaded range (Burger and Gochfeld 2009; Reed et al. 2014; Briceño et al. 2019), as well in our study area, such as peregrine falcon (Sánchez-Mateos and Sánchez-Albarrán 2017) and golden eagle Aquila chrysaetos (E. Navarro pers. comm.). We recorded remains of monk parakeets in several nests of black kites, booted eagles and peregrine falcons and observed their hunting attempts on flying parakeets. The other two raptor species are of similar size and behaviour, and also include birds in their diets (Forsman 1999). Thus, we considered the five raptor species as potential predators. We did not find evidence for other bird species, such as corvids, preying upon monk parakeets or their nests. For storks and raptors, each nest corresponds to a single breeding pair, whereas for parakeets each nest can house from one to several breeding pairs accommodated in different chambers (range: 1–35 active chambers). A parakeet nest was classified as being associated with a stork if they shared the same nesting substrate, if the parakeet nest was located on the same structure (i.e., the same tree or electricity pylon) as the stork nest, or if the parakeet nest was within a radius of 15 m of a stork nest (Fig. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 134 (Fig.1).1). Although mammals and snakes can predate on parakeets, they were not considered in this study as we do not have accurate information about their distribution and abundance. However, their potential impact on parakeets is discussed based on nest substrate (see previous). Interactions between parakeets and raptors We recorded the responses of parakeets towards intruding raptors (i.e., raptors flying within less than 15 m of an active parakeet nest) across the study area, following previous work conducted on a similar species, the rose-ringed parakeet (Psittacula krameri; Hernández-Brito et al. 2014). Parakeet responses were classified as stay (i.e., when parakeets stayed in their nests, showing no sign of fear toward the raptor), mobbing (i.e., when parakeets flew to attack the approaching raptor), or flush (i.e., when parakeets flew away from the raptor). Statistical analyses We used Generalised Linear Models to test if parakeet-stork associations were more likely in rural than in urban areas (logistic link functions, binomial error distributions) by considering all parakeet nests present in the study area and including the habitat where they were located (i.e., urban or rural) as an explanatory variable. We then evaluated if parakeets used rural stork nests (probability of parakeet-stork association considering all stork nests present in the rural areas; logistic link functions, binomial error distributions) based on a combination of conspecific density, type of substrate and predation risk. Conspecific density was obtained as the relative position of each parakeet nest within the parakeet population. We used the formula Σexp(−dij)*A, where dij is the linear distance between each parakeet nest i and all parakeet nests j, and A is the number of chambers per nest j (Moilanen and Hanski 1998). Higher values of this index point to a higher density of conspecifics around a selected location. Predation risk Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 135 was assessed by using the aggregation of raptors nests as a proxy, calculated using the same index explained previously (note that here A always equals 1). Models for the probability of parakeet-stork associations were separately run for 2014 and 2015 because of convergence problems when using generalised linear mixed models and nest as a random term. Finally, we related the probability of nest abandonment by rural parakeets with nest abandonment by the stork (independent variable), considering if colonies occupied in 2014 remained occupied in 2015 (logistic link functions, binomial error distributions). All continuous variables were included in their linear and quadratic forms and standardised before modelling. Model selection was performed using the Akaike Information Criterion corrected for small sample sizes, AICc; (Burnham and Anderson 2002). Within each set of models (which includes the null model), we calculated the ∆AICci (as the difference between the AICc of model i and that of the best model) and the weight (w) of each model. Models within 2 AICc units of the best one were considered as alternatives and used to perform model averaging (MuMIn package). We considered that a given effect received no, weak or strong statistical support when the 95% confidence interval (CI) strongly overlapped with zero, barely overlapped with zero, or did not overlap with zero, respectively. Statistical analyses were conducted in R 3.1.2 (R Core Team, 2013). Differences in behavioural responses of parakeets toward approaching raptors were compared among habitats (urban or rural) and between nests associated or not associated with storks using generalised linear models (multinomial error distribution). We included the number of parakeets present in each raptor intrusion and the interacting raptor species as covariates to control for their potential effects. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 136 RESULTS Protective nest associations We recorded more than 900 monk parakeet nests, most of them located in the urban habitat, while avian predators were much more abundant in rural habitats (Fig. 2; Table 1). Parakeets nesting in rural habitats were rare and mainly associated with storks (97.06% of parakeet nests associated with storks in 2014, n = 34; 73.53% of parakeet nests associated with storks in 2015, n = 34), an association that was near completely absent among urban parakeets (habitat (urban): estimate: -2.30; 95% CI: − 2.88 - -1.73). Importantly, parakeets bred mainly associated with stork nests located in pylons (70 and 100% of associated nests in 2014 and 2015 were in pylons, Table Table1).1). However, models run for 2014 and 2015 show that the probability of parakeet-stork association was only related to habitat (i.e., more likely in rural than in urban areas; Table 2). Fig. 2 Study area. Urban (dark grey) and surrounding rural (white) areas of Madrid Metropolitan area (40° 21′ 03.1“ N, 3° 30’ 06.1” W). Different coloured points show the location of nests of raptors (red), storks (yellow), and parakeets associated (blue) or not associated (purple) with storks. Black dashed lines are rivers Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 137 Table 1 Abundance of species in the study area. Number of nests of Monk parakeets, white storks and avian predators in urban and rural areas. The type of substrate (i.e., pylon, tree or roof) is indicated for Monk parakeet and white stork nests Species 2014 2015 Urban Rural Urban Rural Monk parakeets Myiopsitta monachus 867 34 890 34 Associated with storks Pylons 0 23 0 25 Trees 9 10 9 0 Roofs 0 0 0 0 Not associated with storks Pylons 2 0 2 0 Trees 856 1 879 9 Roofs 0 0 0 0 White storks Ciconia ciconia 41 466 47 440 Pylons 6 233 7 209 Trees 32 217 37 216 Roofs 3 16 3 15 Black kites Milvus migrans 0 239 0 244 Common buzzards Buteo buteo 0 15 0 14 Booted eagles Hieraaetus pennatus 0 5 0 4 Northern goshawks Accipiter gentilis 0 3 0 3 Red kites Milvus milvus 0 1 0 1 Peregrine falcon Falco peregrinus 6 0 6 0 Within rural habitats, stork nests were not used at random, and parakeets selected, among those available, stork nests located in areas where conspecifics were more abundant and predation risk was lower (Table 3; Fig. 3). It is worth noting that the correlation between conspecific density and predation risk increased from 2014 to 2015 (− 0.38 and − 0.48, respectively), so predation risk received weaker support in models obtained for the second year. However, when conspecific density was excluded from models, predation risk was strongly related to stork nests also used in 2015 (estimate: - 1.34, 95% CI: − 2.60 - -0.47). Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 138 ** * 97.5% 1852.17 -3.58 0.03 6577.41 2.5% -1814.86 -5.61 -1.73 -6642.7 Estimate 18.66 -4.59 -0.85 -32.65 Variables substrate (pylon) habitat (urban) year (2015) habitat (urban)*substrate (pylon) Weight 0.58 0.24 0.13 0.06 0 0 0 0 0 0 0 ∆AICc 0 1.81 3.08 4.64 22.49 26.35 26.95 84.12 85.48 399.36 400.26 AICc 234.58 236.4 237.66 239.22 257.07 260.93 261.53 318.7 320.07 633.94 634.84 k 5 4 4 3 4 3 2 3 2 1 2 Model substrate*habitat + year substrate*habitat substrate + habitat + year substrate + habitat habitat*year habitat + year habitat substrate + year substrate null year R 2 =0.70 Table 2 Factors affecting the probability of association between parakeets and storks. Relative importance of habitat (urban and rural) and type of substrate (pylon, tree or roof) on the probability of protective nesting associations between monk parakeets Myiopsitta monachus and white storks Ciconia ciconia. Estimates and 95% confidence intervals (2.5 and 97.5%) were assessed after model averaging (∆AIC ≤2). We considered that a given variable has no, weak or strong support when the 95% confidence interval strongly overlapped zero, barely overlapped zero (*), or did not overlap zero (**), respectively. k: number of parameters. AICc: Akaike Information Criterion corrected for small sample sizes. ∆AICc: difference between the AICc of model i and that of the best-supported model (i.e. the model with the lowest AICc); w: Akaike weights. R 2 : measure of how well the model explains the data Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 139 ** ** ** 97.50% -1.94 3.81 2563.8 2564.79 97.50% 2.84 13256.1 13745.84 1.25 2.50% -5.47 1.56 -2533.35 -2532.36 2.50% 1.19 - 13219.31 - 13742.57 -2.75 Estimate -3.7 2.69 15.22 16.21 Estimate 2.02 18.4 1.63 -0.75 Variables predation risk conspecific density substrate (pylon) substrate (tree) Variables conspecific density substrate (pylon) substrate (tree) predation risk weight 0.67 0.33 0 0 0 0 0 0 weight 0.36 0.31 0.19 0.15 0 0 0 0 ∆AICc 0 1.38 21.19 26.31 66.34 72.52 75.76 95.04 ∆AICc 0 0.32 1.26 1.8 48.71 58.04 63.18 92.88 AICc 107.56 108.94 128.75 133.87 173.9 180.08 183.32 202.61 AICc 101.07 101.39 102.34 102.87 149.78 159.11 164.26 193.96 k 3 5 4 2 4 2 3 1 k 4 2 3 5 4 3 2 1 Model 2014 predation risk + conspecific density predation risk + conspecific density + substrate conspecific density + substrate conspecific density predation risk + substrate predation risk substrate null Model 2015 conspecific density + substrate conspecific density predation risk + conspecific density predation risk + conspecific density + substrate predation risk + substrate substrate predation risk null 2014: R 2 =0.39 2015: R 2 =0.40 Table 3 Factors affecting the probability of association between parakeets and storks in rural areas. Relative importance of predation risk, conspecific aggregation and substrate on the probability of protective nesting associations between Monk parakeets Myiopsitta monachus and white storks Ciconia ciconia. Estimates and 95% confidence intervals (2.5 and 97.5%) were assessed after model averaging (∆AIC ≤2). We considered that a given variable has no, weak or strong support when the 95% confidence interval strongly overlapped zero, barely overlapped zero (*), or did not overlap zero (**), respectively. Models were run separately for 2014 and 2015. k: number of parameters. AICc: Akaike Information Criterion corrected for small sample sizes. ∆AICc: difference between the AICc of model i and that of the best-supported model (i.e. the model with the lowest AICc); w: Akaike weights. R2: measure of how well the model explains the data Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 140 Fig. 3 Probability of parakeet-stork association in rural habitats. Parakeets select rural stork nests located farther from predators (aggregation of predators) and surrounded by larger densities of conspecifics (aggregation of parakeets). Estimates (solid lines), confidence intervals (dashed lines) and raw data (black dots) are shown for 2014 and 2015 Probability of nest abandonment by parakeets From 2014 to 2015, 44% of the 34 rural parakeet nests were abandoned, while all urban parakeet nests remained active. This high rate of abandonment recorded among rural nests was strongly explained by the abandonment of the nest by the stork (Table 4). Substrate, conspecific density and predation risk were weakly supported, with nests located in pylons, far from conspecifics and in areas with a low density of predators being more prone to abandonment when they were abandoned by storks. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 147 inbreeding depression or the evolution of adaptations to the new habitatis an important part of the colonisation process (Sakai et al. 2001; Crooks 2005; Lockwood 2013). Spread lag phases may take years or decades, can vary among species and populations of the same species subject to different ecological conditions (Aagaard 2014) and are highly unpredictable (Coutts et al. 2018). Therefore, it is not prudent to assume that exotic species that have been observed in urban areas for a long period will remain strictly urban in the future. In fact, the distribution of monk parakeets invading Israel has increased and shifted from predominantly urban areas to agricultural landscapes in less than two decades (Coutts et al. 2018). This may have been facilitated by the longterm decline and poor conservation status of raptors in that country (Yom-Tov et al. 2012). Thus, accepting invasive species as part of the urban ecosystem may sometimes result in their spread into adjacent rural landscapes, where they could have different, often unknown impacts (Botham et al. 2009, Moreira-Arce et al. 2014), such as crop damage in the case of monk parakeets (Senar et al. 2016; Postigo et al. 2017). The monk parakeet was first introduced in Spain in 1976, increasing its distribution since then at a rate of 8.14 grid cells (5 × 5 km) per year (Abellán et al. 2017). However, it was not until recently that the Spanish population began to grow exponentially, increasing from c. 6000 individuals in 2010 to c. 20,000 in 2015 distributed across > 130 urban populations (Molina et al. 2016). Therefore, it seems that the species has overcome the lag phase, and the protective nesting association facilitating its spread outside of cities, rather than anecdotal, may be occurring in other areas. Predictive models indicate that there is still plenty of suitable habitat for the species (Muñoz et al. 2006; Molina et al. 2016). Furthermore, white storks are widely distributed across Spain (with > 33,000 nests in 2004; Molina et al. 2004) and populations are rapidly increasing thanks to the use of human-related food subsidies Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 148 (invasive American crayfish and rubbish dumps (Blanco 1996; Tablado et al. 2010; Sanz-Aguilar et al. 2015). In fact, in recent years, monk parakeets also spread from another Spanish city (Zaragoza, 270 km distant to Madrid), by nesting in stork nests (J.L. Tella obs. Pers.), but such a spread was halted by the responsible authorities by shooting the whole parakeet population. Therefore, the risk of monk parakeet expansion from urban habitats, thanks to the widespread distribution of white storks, should be considered when designing management strategies for this highly invasive species, which is growing exponentially in Mediterranean countries (Postigo et al. 2019), including Spain, Portugal, France, Italy, Greece, Morocco, and Israel, where the two species coexist (Hagemeijer and Blair 1997; Postigo et al. 2019). Management actions could be required in the case rural populations of monk parakeets would cause significant impacts (Postigo et al. 2019), although these actions usually show low support from the society (Shackleton et al. 2019), even lower when dealing with charismatic species such as parakeets (Crowley et al. 2019). Consequently, more research and awareness campaigns are necessary not only to know the actual magnitude of the impacts derived from invasive species (Blackburn et al. 2010) but also to make management actions effective. In our case, actions only focus on the avoidance of monk parakeets nesting in white stork nests may not be efficient. On the one hand, antinesting devices installed in pylons for white storks do not prevent their nesting, even after great management efforts (Maricato et al. 2016). On the other hand, nest removal would be not efficient because both species show strong fidelity to their nesting substrates and often rebuild their nests very soon (Sol et al. 1997; Vergara et al. 2006; Pruett-Jones et al. 2007; Burger and Gochfeld 2009). Contrarily, actions aimed to improve predator populations as biological controllers, should be effective to halt the spread of this species into rural areas in the long term. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 149 This study assesses the nesting association between an invasive bird, the monk parakeet, and a native bird species, the white stork, showing a commensalism relationship in which parakeet colonies associated with stork nests benefit from the effective antipredatory defence of storks. This association was more likely in rural areas, where predation pressure is higher than in nearby urban ones, assisting thus the spread of monk parakeets across the rural environment. Moreover, the abandonment of parakeet colonies after the previous nest abandonment of associated storks, as well as their different behavioural reactions against raptors when associated with storks, suggest that parakeets have a strong dependence of their hosts. However, this protective association is limited as parakeet colonies also avoided high densities of breeding raptors in the study area. Without the facilitation provided by storks, the biotic resistance from the raptor community prevents the invasion success of parakeets. Future studies are needed to assess the complexity of interactions between invasive species and the recipient community, which may be fundamental to develop effective management plans against biological invasions. ACKNOWLEDGEMENTS We thank J. López for helping during fieldwork and the owners of Caserío del Henares and El Piul for allowing us to access their private lands (the rest of the study area was access-free). Ó. Frías and F. Martínez also helped in the fieldwork and contributed data on the location of nests of raptors, storks and parakeets. Logistic and technical support were provided by Doñana ICTS-RBD. This study was supported by the Severo Ochoa Program (SVP-2014-068732) and Action COST ES1304 (ParrotNet). Logistical and technical support for fieldwork was provided by Doñana ICTS-RBD. Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Section 2: Chapter V 150 REFERENCES − Aagaard, K., & Lockwood, J. (2014). Exotic birds show lags in population growth. 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Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Supplementary Material Section 3-Chapter VIII 338 Pyrrhura cruentata Neotropical 1 0 0 Pyrrhura frontalis Neotropical 0 0 1 Pyrrhura griseipectus Neotropical 1 0 0 Pyrrhura molinae Neotropical 1 0 1 Pyrrhura perlata Neotropical 1 0 0 Thectocercus acuticaudatus Neotropical 0 1 1 Trichoglossus haematodus Australasia 0 0 1 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Supplementary Material Section 3-Chapter VIII 339 Table S2. List of plant species where parrots wasted food. We show the species, family and the number of times we detected food wasting at the species. Species Family Number of observations Acacia angustissima Fabaceae 16 Acacia aroma Fabaceae 1 Acacia auriculiformis Fabaceae 12 Acacia karroo Fabaceae 13 Acacia mearnsii Fabaceae 10 Acacia melanoxylon Fabaceae 11 Acaena splendens Rosaceae 1 Acrocomia totai Arecaceae 65 Aesculus hippocastanum Sapindaceae 1 Agonandra brasiliensis Opiliaceae 2 Ailanthus altissima Simaroubaceae 2 Aiphanes minima Arecaceae 2 Aizoon canariense Aizoaceae 1 Albizia carbonara Fabaceae 1 Albizia guachapele Fabaceae 2 Albizia julibrissin Fabaceae 1 Albizia procera Fabaceae 9 Albizia saman Fabaceae 19 Anacardium excelsum Anacardiaceae 3 Anacardium occidentale Anacardiaceae 9 Anadenathera colubrina Fabaceae 2 Anisocapparis speciosa Capparidaceae 4 Annona muricata Annonaceae 2 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Supplementary Material Section 3-Chapter VIII 340 Arachis hypogaea Fabaceae 36 Araucaria angustifolia Araucariaceae 303 Araucaria araucana Araucariaceae 34 Araucaria bidwillii Araucariaceae 51 Araucaria cunninghamii Araucariaceae 5 Arbutus unedo Ericaceae 7 Ardisia escallonioides Primulaceae 4 Asparagus arborescens Asparagaceae 2 Aspidosperma quebracho-blanco Apocynaceae 2 Astronium urundeuva Anacardiaceae 1 Attalea butyracea Arecaceae 2 Attalea phalerata Arecaceae 0 Attalea princeps Arecaceae 3 Attalea speciosa Arecaceae 9 Attalea totai Arecaceae 1 Averrhoa carambola Oxalidaceae 3 Azadirachta indica Meliaceae 2 Bactris gasipaes Arecaceae 1 Bambusa vulgaris Poaceae 4 Betula pendula Betulaceae 4 Brachychiton populneus Malvaceae 3 Brosimum alicastrum Moraceae 1 Bucida buceras Combretaceae 4 Bursera simaruba Burseraceae 3 Bursera tomentosa Burseraceae 2 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Supplementary Material Section 3-Chapter VIII 341 Byrsonima crassifolia Malpighiaceae 7 Byrsonima pachyphylla Malpighiaceae 34 Byrsonima verbascifolia Malpighiaceae 5 Cajanus cajan Fabaceae 1 Callicarpa ampla Lamiaceae 3 Calophyllum antillarum Calophyllaceae 6 Carapa guianensis Meliaceae 2 Carduus thoermeri Asteraceae 112 Carya illinoinensis Juglandaceae 7 Cassia javanica Fabaceae 2 Castanea sativa Fagaceae 3 Castilla elastica Moraceae 1 Casuarina cristata Casuarinaceae 1 Casuarina equisetifolia Casuarinaceae 11 Catalpa bignonioides Bignoniaceae 7 Cecropia peltata Urticaceae 1 Cecropia schreberiana Urticaceae 9 Cedrela odorata Meliaceae 15 Cedrus atlantica Pinaceae 1 Cedrus libani Pinaceae 1 Ceiba pentandra Malvaceae 28 Ceiba penthandra Malvaceae 1 Ceiba speciosa Malvaceae 9 Celtis australis Ulmaceae 45 Centaurea solstitialis Asteraceae 46 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Supplementary Material Section 3-Chapter VIII 342 Cerastium glomeratum Caryophyllaceae 3 Ceratonia siliqua Fabaceae 5 Cercis siliquastrum Fabaceae 67 Chenopodium murale Chenopodiaceae 4 Cipadessa baccifera Meliaceae 1 Citharexylum donnell-smithii Verbenaceae 1 Citharexylum spinosum Verbenaceae 7 Citrus × aurantium Rutaceae 2 Citrus × sinensis Rutaceae 37 Citrus aurantifolia Rutaceae 1 Clusia pratensis Clusiaceae 1 Cnidoscolus quercifolius Euphorbiaceae 1 Coccoloba caracasana Polygonaceae 1 Coccoloba uvifera Polygonaceae 52 Cocos nucifera Arecaceae 7 Coffea arabica Rubiaceae 5 Commiphora leptophloeos Burseraceae 1 Compsoneura sprucei Myristicaceae 1 Condalia buxifolia Rhamnaceae 8 Cordia alba Boraginaceae 1 Cordia collococca Boraginaceae 5 Cordia cymosa Boraginaceae 1 Corymbia clarksoniana Myrtaceae 7 Corymbia tessallaris Myrtaceae 2 Couepia grandiflora Chrysobalanaceae 15 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Supplementary Material Section 3-Chapter VIII 343 Couepia polyandra Chrysobalanaceae 1 Couma macrocarpa Apocynaceae 2 Crataegus monogina Rosaceae 1 Croton magdalenensis Euphorbiaceae 1 Cupania americana Sapindaceae 3 Cupressus sempervirens Cupressaceae 87 Curatella americana Dilleniaceae 16 Cyanophalla retusa Capparidaceae 2 Cyperaceus sp. Cyperaceae 1 Cyttaria harioti Cyttariaceae 2 Delonix regia Fabaceae 15 Dendropanax latilobus Araliaceae 1 Dovyalis caffra Salicaceae 47 Dracaena draco Asparagaceae 2 Drimys winteri Winteraceae 1 Elaeis guianensis Arecaceae 1 Emmotum nitens Icacinaeae 7 Eriobotrya japonica Rosaceae 75 Erythrina caffra Fabaceae 21 Erythrina fusca Fabaceae 16 Erythrina poeppigiana Fabaceae 10 Erythroxylum deciduum Erythroxylaceae 6 Eucalyptus camaldulensis Myrtaceae 45 Eucalyptus globulus Myrtaceae 35 Eucalyptus miniata Myrtaceae 5 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Supplementary Material Section 3-Chapter VIII 344 Eucryphia cordifolia Cunoniaceae 1 Eugenia uniflora Myrtaceae 38 Ficus benghalensis Moraceae 19 Ficus carica Moraceae 115 Ficus insipida Moraceae 2 Ficus lyrata Moraceae 27 Ficus macrophylla Moraceae 57 Ficus maxima Moraceae 1 Ficus microcarpa Moraceae 106 Ficus nymphaeifolia Moraceae 1 Ficus religiosa Moraceae 9 Ficus rubiginosa Moraceae 16 Ficus sp. Moraceae 1 Ficus stuhlmannii Moraceae 6 Ficus trigonata Moraceae 2 Flacourtia inermis Salicaceae 5 Fraxinus angustigolia Oleaceae 33 Geoffroea decorticans Fabaceae 1 Gleditsia triacanthos Fabaceae 18 Gliricidia sepium Fabaceae 9 Gmelina arborea Lamiaceae 15 Grevillea glauca Proteaceae 3 Grevillea robusta Proteaceae 91 Guarea guidonia Meliaceae 27 Guazuma ulmifolia Malvaceae 1 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Supplementary Material Section 3-Chapter VIII 345 Hancornia speciosa Apocynaceae 3 Handroantus chrysanthus Bignoniaceae 2 Helianthus annuus Asteraceae 99 Heliocarpus sp. Malvaceae 1 Hibiscus rosa-sinensis Malvaceae 4 Hirtella rugosa Chrysobalanaceae 2 Hymenaea courbaril Fabaceae 1 Hymenaea stigonocarpa Fabaceae 17 Hypochaeris radicata Asteraceae 2 Inga adenophylla Fabaceae 1 Inga chocoensis Fabaceae 1 Inga cylindrica Fabaceae 3 Inga edulis Fabaceae 1 Inga feuilleei Fabaceae 1 Inga punctata Fabaceae 1 Inga spectabilis Fabaceae 11 Jacaranda mimosifolia Bignoniaceae 39 Jatropha hieronymi Euphorbiaceae 11 Jatropha humboldtiana Euphorbiaceae 7 Jatropha mollisima Euphorbiaceae 4 Juglans regia Juglandaceae 3 Juniperus procera Cupressaceae 1 Juniperus virginiana Cupressaceae 9 Lagerstroemia indica Lythraceae 2 Lagerstroemia speciosa Lythraceae 9 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Supplementary Material Section 3-Chapter VIII 346 Laureliopsis philippiana Atherospermataceae 2 Leucaena leucocephala Fabaceae 2 Ligustrum japonicum Oleaceae 38 Ligustrum lucidum Oleaceae 13 Livistona chinensis Arecaceae 165 Luehea seemannii Malvaceae 4 Magonia pubescens Sapindaceae 2 Malus domestica Rosaceae 17 Mangifer indica Anacardiaceae 96 Mangifera zeylanica Anacardiaceae 1 Manilkara bidentata Sapotaceae 3 Maprounea guianensis Euphorbiaceae 2 Maranthes panamensis Chrysobalanaceae 7 Mauritia flexuosa Arecaceae 46 Mauritiella armata Arecaceae 1 Maytenus boaria Celastraceae 5 Melaleuca citrina Myrtaceae 17 Melia azedarach Meliaceae 311 Melicoccus bijugatus Sapindaceae 10 Miconia impetiolaris Melastomataceae 10 Misodendrum linearifolium Misodendraceae 2 Morus alba Moraceae 84 Morus nigra Moraceae 127 Mouriri pusa Melastomataceae 2 Musa × paradisiaca Poaceae 3 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Supplementary Material Section 3-Chapter VIII 347 Myrceugenia exsucca Myrtaceae 1 Myrtus communis Myrtaceae 16 Neea theifera Nyctaginaceae 16 Nephelium lappaceum Sapindaceae 1 Nothofagus dombeyi Nothofagaceae 3 Nothofagus obliqua Nothofagaceae 10 Ochroma pyramidale Malvaceae 1 Olea europaea Oleaceae 49 Olea europaea sylvestris Oleaceae 21 Onopordum acanthium Asteraceae 1 Oryza sativa Poaceae 11 Pachira quinata Malvaceae 10 Parkia biglandulosa Fabaceae 4 Parkinsonia aculeata Fabaceae 3 Parkinsonia praecox Fabaceae 1 Parodiodendron marginivillosum Picrodendraceae 2 Passiflora edulis Passifloraceae 2 Patellifolia patellaris Amaranthaceae 5 Patellifolia procumbens Amaranthaceae 1 Peltophorum pterocarpum Fabaceae 1 Persea americana Lauraceae 11 Phalaris canariensis Poaceae 3 Phoenix canariensis Arecaceae 260 Phoenix dactylifera Arecaceae 437 Phytolacca dioica Phytolaccaceae 33 Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda Código seguro de Verificación : GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda | Puede verificar la integridad de este documento en la siguiente dirección : https://sede.administracionespublicas.gob.es/valida ÁMBITOPREFIJO CSV FECHA Y HORA DEL DOCUMENTO GEISER GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda 30/10/2020 13:07:44 Horario peninsular Nº registro DIRECCIÓN DE VALIDACIÓN O00008744e2000054554 https://sede.administracionespublicas.gob.es/valida GEISER-46bd-7fd7-a9cf-43bd-b44e-b9e3-7a56-3fda