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Are invasive House Sparrows a nuisance for native avifauna when scarce?

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Michelle García-Arroyo1, Diego Santiago-Alarcón2, Javier Quesada3 & Ian MacGregor-Fors1

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1 Red de Ambiente y Sustentabilidad, Instituto de Ecología, A. C., Carretera antigua a Coatepec
351, El Haya, 91073 Xalapa, Veracruz, Mexico

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2 Red de Ecología y Conservación de Vertebrados, Instituto de

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Ecología, A. C., Carretera antigua a Coatepec 351, El Haya,

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91073 Xalapa, Veracruz, Mexico

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3 Department of Vertebrates, Natural Sciences Museum of Barcelona, Parc de la Ciutadella s/n,
08003 Barcelona, Catalonia, Spain

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Correspondenc author: Ian MacGregor-Fors

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ian.macgregor@inecol.mx; macgregor.ian@gmail.com

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Abstract

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Biological invasions are the second most important cause of species extinction. Aided by processes

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such as transportation and urbanization, exotic species can establish and spread to new locations,

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causing changes in the function and structure of ecosystems. The House Sparrow is a widespread

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and highly abundant landbird associated to human presence. Previous studies performed in

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urban landscapes have suggested that this species could be acting, in synergy with urbanization,

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as a potential threat to native urban avian assemblages. In this study we assessed the relationship

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between House Sparrow density and native bird species richness in a region where the sparrows

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are scarce and sparsely distributed. We surveyed bird assemblages in and around four small-sized

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human settlements, considering three conditions in relation to House Sparrow presence: urban

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invaded, urban non-invaded, and non-urban non-invaded. To assess the potential detrimental

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role of House Sparrows on native bird species richness, we measured, additionally to sparrow

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densities, 20 predictor variables that describe vegetation structure and complexity, as well as

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urban infrastructure and human activities across four seasons of 1 year. Our results show that

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maximum shrub height was positively related to bird species richness, built cover was negatively

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associated with it, and House Sparrow invaded sites were related to a significant decrease of bird

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species richness, with increasing richness loss when more sparrows were present. Thus, we here

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provide evidence that urban areas can act in synergy with the presence of House Sparrows (even in

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low densities) in the urban-related species richness decline pattern.

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Keywords:

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Biological invasions, Bird density, Passer domesticus, Species composition, Species richness,

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Urban ecology

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Introduction

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Biological invasions are considered one of the main drivers of species extinctions, altering species

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richness and composition of native communities at different spatiotemporal scales (Bellard et al.

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2016). When the individuals of exotic species establish and colonize new locations, successful

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biological invasions occur (Blackburn et al. 2011) and may alter local environmental processes

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and the structure of local native communities (e.g., nutrient cycles, trophic networks, fire and

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erosion regimes; Pyšek et al. 2012; Ricciardi et al. 2013; Simberloff et al. 2013). Although invasive

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birds are abundant across the globe (Blackburn et al. 2009), the magnitude and variability of

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their impact on native assemblages remains poorly understood (Kumschick and Nentwig 2010). It

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is notable that three avian species have been included in the list of 100 worst invasive alien

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species (Lowe et al. 2000; but see Kumschick et al. 2016): Common Myna (Acridotheres tristis),

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Red-vented Bulbul (Pycnonotus cafer; but see Thibault et al.2018), and European Starling

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(Sturnus vulgaris). These three species alone have been responsible for massive damages to

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crops and infrastructure, but also for spreading diseases, and displacing native avifauna through

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predation and competition for nest cavities (Fisher and Wiebe 2005; Harper et al. 2005; Tindall et

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al. 2007; Grarock et al. 2012).

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Cities are key components for avian invasions, not only as hubs for the deliberate trading of pets,

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but also by promoting the establishment and spread of diverse bird species in highly predictable

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systems (Vitousek et al. 1997; Sax and Brown 2000; Shochat 2004; Shochat et al.2010). The

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filtering of regional avifaunas in urban settings generally results in depauperate avian

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assemblages, especially in heavily urbanized conditions, a niche that has been heavily exploited

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by generalists, often exotic and/or invasive species (Chace and Walsh 2006; Aronson et al. 2014;

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La Sorte et al. 2018). Given that many of these generalist urban exploiters are prone to

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experience population explosions in urban areas, they frequently dominate urban bird

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assemblages (Sol et al. 2014).

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Urban invasive birds have been accounted for economic losses due to damages to buildings and

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other urban structures (Pimentel et al. 2001, 2005; Booy et al. 2017), as well as the spread of

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diseases on a global scale (Pedersen et al. 2006). However, there is a lack of agreement on the

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ecological impacts that invasive birds pose on native species (Linz et al.2007; Strubbe and

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Matthysen 2007; MacGregor-Fors et al.2010, 2011; Mori et al. 2017; González-Oreja et al. 2018;

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Luna et al. 2018). One of the most widespread urban-related invasive bird species is the House

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Sparrow (Passer domesticus), a species considered to be native to Eurasia and North Africa and

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that has been associated with humans for 10,000 years, since the appearance of agricultural

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practices (Anderson 2006; Sætre et al. 2012). This sparrow has been either intentionally or

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unintentionally introduced by humans in Australia, New Zealand, North America, South America,

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and South Africa (Anderson 2006). Regarding its North American invasion, it was successfully

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introduced to Northeastern United States in the 1850s and arrived to Mexico around the 1910s,

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establishing numerous and dense populations that expanded across the country in following

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decades, reaching Mexico City by 1930 (Wagner 1959). House Sparrow populations resulting

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from these invasion events have continued their range expansion southward to Central America

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(Anderson 2006).

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House Sparrows are ecologically and physiologically plastic, with an extensive array of nesting

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habits (Kimball 1997; Nhlane 2000; Peach et al. 2008; Hoi et al. 2011), foraging behaviors, and

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dietary breadth (Guillory and Deshotels 1981; Kalmus 1984; Flux and Thompson 1986; Anderson

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2006). Although its main food sources are seeds, it has an omnivorous diet in urban

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environments, ranging from nectar, fruits, insects, and even discarded human food leftovers

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(Stidolph 1974; Gavett and Wakeley 1986; Clergeau 1990; Moulton and Ferris 1991; Leveau

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2008; MacGregor-Fors et al. 2020). Behaviorally, the House Sparrow is aggressive with both its

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conspecifics and heterospecifics, often competing for nesting cavities and food resources

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(Kalinoski 1975; Gowaty 1984; Radunzel et al. 1997; Anderson 2006). It is also known to be an

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important source of pathogens (Rappole and Hubálek 2003; e.g., avian pox and malaria, West

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Nile Virus; Anderson 2006; Delgado-V and French 2012). Albeit the undeniable success of House

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Sparrows in North America, population declines have been recorded in the past decades along

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urban-agricultural landscapes of Western Europe (Summers-Smith 2003).

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Previous studies have shown negative relationships between the presence and abundance of

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House Sparrows and other native land birds. For instance, in a Central Western Mexico

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medium-size city, avian assemblages dominated by House Sparrows had lower bird species

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richness (MacGregor-Fors et al. 2010). In another study performed in Mexico City, the

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abundance of some native bird species showed to be negatively related with the presence and

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abundance of House Sparrows (i.e., Berylline Hummingbird–Amazilia beryllina, Black-headed

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Grosbeak–Pheucticus melanocephalus), with lower average abundance per point count ranging

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from 40% to 300% decreases (Ortega-Álvarez and MacGregor-Fors 2010). Moreover, the

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abundance of rare native birds was negatively associated with sites used by House Sparrows for

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roosting and breeding, such as lamp poles in a west-central Mexican city (MacGregor-Fors and

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Schondube 2011). Yet, results of a recent study performed in urban greenspaces of three

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Mexican cities suggest that House Sparrows are not related with declines in native species

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richness (González-Oreja et al. 2018). Based on all of the above, we consider that there is

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enough correlative evidence to acknowledge that House Sparrows can represent a potential

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competitor able to displace native species (Schondube et al. 2009).

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In this study we assessed the relationship between House Sparrow density and native bird

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species richness in scenarios where sparrows are scarce and sparsely distributed. It is notable

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that these conditions, where House Sparrows are not hyper-abundant differ to those of

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previous studies focused on the potential effects to native avifauna, where sparrow densities

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are high (MacGregor-Fors et al. 2010; Ortega-Álvarez and MacGregor-Fors 2010). Thus, we

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surveyed bird assemblages in and around four small-sized human settlements in Central

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Veracruz (Mexico), where House Sparrows are present in low numbers, considering three

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different conditions: urban invaded, urban non-invaded, and non-urban non-invaded. Based on

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contrasting results related to the potential negative relationship between House Sparrows and

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native bird species richness, we tested the following hypotheses: (1) low densities of House

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Sparrows are associated with a lower bird species richness and composition, holding the

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pattern of previous studies evidencing the negative relationship regardless of sparrows’

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densities, and (2) low densities of House Sparrows do not relate to bird species richness nor its

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composition, and thus do not represent a nuisance for native avifauna when present in low

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densities.

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Methods

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Study area

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We conducted this study in four human settlements from Central Veracruz: Xico, Teocelo, San

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Marcos de León, and Colonia Úrsulo Galván (referred to as Xico, Teocelo, San Marcos, and Úrsulo

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Galván hereafter; Table 1). The largest settlement in the region is Xico, with an extension of 2

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km2 and a population of ~18,650 inhabitants (INEGI 2010), followed by Teocelo (1 km2, ~9950

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inhabitants; INEGI 2010), San Marcos (0.7 km2, ~7250 inhabitants; INEGI 2010), and Úrsulo

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Galván (0.14 km2, ~1700 inhabitants; INEGI 2010). The studied settlements have similar urban in-

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frastructure (mainly composed of one to two story houses, few commercial areas, few buildings

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with over four stories) and are embedded in a landscape with similar characteristics (hilly

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topography, presence of multiple water streams, and similar climate; INAFED 2010). It is notable

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that the study region was originally covered, in general, by tropical montane cloud forest, which

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has been partially replaced over the last century by shade coffee plantations, cattle ranches, and

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urban centers (Williams-Linera 2007; García-Franco et al. 2008).

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Study design and field surveys

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We followed a survey design that allowed us to assess the relationship between the presence

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and abundance of House Sparrows and native bird species richness, considering two

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dichotomies: (1) House Sparrow invaded / House Sparrow non-invaded sites and (2) built up

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environments (referred to as urban hereafter) / non-built sites (sensu MacGregor-Fors2010).

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Given that House Sparrows are absent outside urban areas in the region, we considered three

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survey conditions: (1) urban House Sparrow invaded (UI), (2) urban House Sparrow non-invaded

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(UNI), and (3) non-urban House Sparrow noninvaded (NUNI). Due to differing sizes of the studied

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settlements and the presence and distribution of House Sparrows within them, our design was

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unbalanced, with a total of 110 survey sites (Table 1, Fig. 1).

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MG-A performed 5-min point counts (25 m limited radius) from sunrise to 11:00 h, recording all

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birds seen or heard at each survey site in four seasons: spring, summer, fall, and winter (i.e., April

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2016, July 2016, October 2016, January 2017). MG-A measured the exact distance from point-

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count locations to each recorded bird individual with a rangefinder (Bushnell Yardage Pro Sport

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450). We established point counts at least 150 m apart from each other to be considered as

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independent sampling units (Ralph et al. 1996; Bibby et al. 2000; Huff et al. 2000).

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Predictor variables

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We measured 20 predictor variables within the same 25 m radius area in which birds were

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counted, once every surveyed season, to describe the environmental characteristics of each

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survey site. To describe vegetation structure and complexity, we recorded: (1) tree richness

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(morphospecies), (2) tree cover (%), (3) number of trees, (4) maximum tree height (m), (5)

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maximum diameter at breast height of trees (DBH) (cm), (6) shrub richness (morphospecies),

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(7) shrub cover (%), (8) maximum shrub height (m), (9) herbaceous plant richness

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(morphospecies), (10) herbaceous plant cover (%), and (11) maximum herbaceous plant height

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(m). To describe urban infrastructure and human activities, we recorded: (1) number of

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buildings, (2) maximum building height (m), (3) minimum building height (m), (4) number of

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light and electric poles, (5) number of cables, (6) number of windows, (7) passing cars per

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minute, and (8) number of pedestrians per minute. Additionally, we quantified built cover (%)

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in the 25 m radius survey area using satellite images from 2016 on Google Earth Pro (2018).

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Data analysis

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We computed the statistical expectation of species richness for each condition using

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rarefaction procedures with EstimateS, which allows statistical comparisons among treatments

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through the repeated re-sampling of all pooled samples based on their recorded abundances

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(Gotelli and Colwell 2001; Colwell 2013). For comparisons among conditions we contrasted the

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84% confidence intervals of the computed statistical expectations and considered statistical

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differences with α = 0.05 when confidence intervals did not overlap (following MacGregor-Fors

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and Payton 2013). We used 84% confidence intervals as 95% confidence intervals fail to

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indicate statistical differences with α = 0.05 (MacGregor-Fors and Payton 2013). Given that

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sampling effort varied among conditions, we used a factor of extrapolation of 2.5 for the

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smallest sample (i.e., UI) to robustly contrast its species richness calculations with the other

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two conditions at the same sampling effort (i.e., UNI, NUNI) (Gotelli and Colwell 2001; Colwell

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2013).

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We performed a multivariate Bray-Curtis cluster analysis (i.e., average linkage) using the

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package ‘vegan’ in R (Oksanen et al. 2016; R Development Core Team 2018) to describe

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similarities in bird assemblage composition among the studied conditions. Taking into account

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the 20 measured predictor variables and to avoid statistical issues related with multi-collinearity,

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we identified moderate-to-highly correlated variables (i.e., r > 0.5, P < 0.05), keeping those with

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highest variance. We used the remaining variables, including House Sparrow abundance per

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point count, in a generalized additive model (GAM) to explore their relationship with bird species

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richness. We used a GAM given that, as a variant of generalized linear models, additive models

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have different error structures and link functions able to provide a better fit for different types of

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variables, also allowing the use of non-parametric ‘smoothers’ (fitting procedure where the form

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of the curve is not predetermined but estimated through data; Wang 2014) to describe non-

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linear relationships (Crawley 2013). If House Sparrow abundances showed a significant

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relationship with species richness, we conducted a t-test to assess differences in built cover

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between sites with and without House Sparrow records.

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To allow comparisons with results of previous studies in Mexico, we report the number of

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House Sparrows per point count, as well as estimated distance-corrected House Sparrow

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densities by season using Distance 6.2 (Thomas et al. 2010). Distance computes densities

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(ind/ha) based on the detection probability of individuals at increasing distances from the

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observer, as well as standardizing detection rates along concentric surveyed areas (Buckland et

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al. 2001).

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Results

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Over the course of four seasons (i.e., spring, summer, fall, winter) we recorded a total of 89 bird

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species of 29 families (Table S1 in Online Resource 1), of which 55% were recorded uniquely at

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the NUNI condition. In particular, we recorded 84 bird species at the NUNI condition, 36 at the

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UNI condition, and 20 at the UI condition. Nearly 25% of the recorded species are reported in the

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literature to be associated with wellvegetated areas, all of which we recorded at the NUNI

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condition, one of them also recorded at the UNI condition (i.e., Black-throated Green Warbler–

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Setophaga virens), and two at the UI condition (i.e., Magnolia Warbler–Setophaga magnolia,

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Rusty Sparrow–Aimophila rufescens) (Table S1 in Online Resource 1). Bird species richness at the

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UI condition was significantly lower when compared to that of the NUNI condition during almost

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all the year (summer, fall, winter) and compared to the UNI condition during summer (Table 2).

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Regarding species composition, the cluster analysis revealed that the UI condition shared less

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species with UNI and NUNI conditions, thus having a different assemblage composition across

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seasons (β = 0.13; Fig. 2).

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Results of the GAM show that bird species richness was significantly related with season

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(Table 3). After taking into account the smoothing adjustment for the numerical variables (i.e.,

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shrub richness, maximum shrub height, built cover, House Sparrow abundances, passing cars per

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minute), we identified that the relationship between maximum shrub height and bird species

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richness was positive (Fig. 3a), the one with built cover was negative (Fig. 3b), and the one with

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House Sparrow abundances showed three different scenarios (i.e., 0 individuals, 1–5 individuals,

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6–12 individuals; Fig. 3c). Due to the complexity of the interpretation of such trichotomy, we

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calculated the statistical expectation of bird species richness for each scenario, finding a

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significant decrease in bird species richness as the number of House Sparrows increased (Fig. 3c).

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It is notable that we did not find differences for built cover values in sites with and without

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House Sparrow records (t25 = −0.77, p = 0.45; Fig. 4), showing that such decrease in species

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richness was not given by urbanization intensity.

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The number of House Sparrows per point count was of 0.6 individuals during spring, 0.49 in

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summer, 0.45 in fall, and 0.4 in winter. Regarding distance-corrected densities, we recorded the

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highest House Sparrow density during winter (12.6 ind/ha 84% CI: 3.5–45.4), followed by spring

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(5.4 ind/ha 84% CI:2.8–10.4), summer (2.5 ind/ha 84% CI: 1.2–4.8) and fall (2.3 ind/ha 84% CI:

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1.0–5.0).

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Discussion

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The House Sparrow is a widespread and highly abundant landbird associated to humans

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(Aronson et al. 2014; Sol et al. 2014) that could be acting in synergy with urbanization as a

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potential threat to native avian assemblages, even when present in low numbers (MacGregor-

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Fors et al. 2010; Loss et al. 2015). Results of this study showed that vegetation elements are

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positively associated with bird species richness, meanwhile heavily urbanized areas are

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negatively related to it. Furthermore, sites with House Sparrows presence had lower bird species

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richness than non-invaded and non-urban areas. Also, the assemblages of invaded urbanized

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areas were more similar among themselves compared to those of noninvaded and non-urban

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areas. Altogether, our findings suggest the existence of different dynamics among bird species

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within urban areas where invasive sparrows are present, having an effect on both the number

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and composition of bird species.

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Seasonality was related to an increase in bird species richness given by the amount of

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Neotropical-Nearctic migrants recorded in winter. It is noteworthy that our study area is located

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within one of the most important Neotropical Nearctic bird migration routes (Ruelas-Inzunza et

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al. 2005). The positive relationship between maximum height of shrubs and bird species richness

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agrees with previous studies assessing avian ecology along urban-agricultural landscapes

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(Ortega-Álvarez and MacGregor-Fors 2009; Faggi and Caula 2017). This variable, as proxy of

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vegetation at each site, highlights the importance of structural stratification of vegetation for

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birds both in non-urban and urban areas (Cueto and de Casenave 1999; Napoletano et al. 2017).

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Built cover was negatively associated with bird species richness, which also agrees with previous

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studies assessing avian assemblages in cities (MacGregor-Fors and Schondube 2011; Luck et al.

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2013; Schneider and Miller 2014; Faggi and Caula 2017). Actually, this relationship was not

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surprising, as urbanization has been directly linked to a decrease in bird species richness due to

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the loss of a wide variety of food resources, breeding sites, and additional factors inherent to

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urbanization (e.g., cat predation, window collision, parasitism; Santiago-Alarcon and Delgado-V

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2017), among other causes (Emlen 1974; Chace and Walsh 2006).

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Finally, House Sparrow numbers had a gradual negative effect on bird species richness,

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where sites having no sparrows (NUNI, UNI) showing significantly more bird species compared

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to sites with sparrows. Specifically, urban invaded areas (UI), with 1–5 House Sparrows had

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significantly more bird species than sites with 6–12 House Sparrows (Fig. 3). It is important to

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highlight that significant differences in bird species richness in sites where we recorded 1–5 and

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6–12 House Sparrows were not related to built cover, as urbanized sites (invaded and non-

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invaded) had similar values (Fig. 4). Given that a possible confounding factor of the recorded

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relationship between House Sparrows and bird species richness could be the potential

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association with the presence and abundance of other urban-related species (i.e., Great-tailed

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Grackle– Quiscalus mexicanus, Rock Pigeon–Columba livia, Tropical Kingbird–Tyrannus

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melancholicus), we assessed potential correlations between the presence and abundance of the

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most frequently recorded species with House Sparrows data. However, we found no significant

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or strong correlations between House Sparrow abundance and the abundance of other

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common urban-associated species (rS ≤ |0.13|, p-values <0.53; Table S2 in Online Resource 1).

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Therefore, our conclusion regarding the negative relationship between House Sparrows and

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native bird species richness holds true.

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Altogether, our results add information to the scarce evidence that this invasive sparrow could

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be acting as a driver of native urban bird assemblages, even when present in low densities. It is

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important to note that House Sparrow numbers recorded in this study were much lower (i.e.,

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10–32 times lower in terms of relative abundance and 1.6–3 times lower in terms of density)

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than those reported in previous studies (i.e., ~20 ind/point count in MacGregor-Fors et al. 2010,

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9.5–33.3 ind/ha in MacGregor-Fors et al. 2017; ~7 ind/point count in Ortega-Álvarez and

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MacGregor-Fors 2011a). Yet, similar low densities are reported for some of the native

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populations of the House Sparrow (Šálek et al. 2015), where this species is considered at risk

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(Summers-Smith 2003; BirdLife International 2004; Shaw et al. 2008).

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Previous evidence has suggested that not only House Sparrows could represent a threat to

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similar sized and smaller granivore species through direct antagonistic interactions (Schondube

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et al. 2009), but also to species from other guilds and sizes, such as hummingbirds (Ortega-

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Álvarez and MacGregor-Fors 2010), as well as species with similar nesting habits (e.g., bluebirds,

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swallows; Kalinoski 1975; pers. obs.). House Sparrow presence along with the threats of

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urbanization (e.g., introduced predators, pollution, habitat destruction; Santiago-Alarcon and

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Delgado-V 2017) and indirect interactions (Marzal et al. 2011, e.g., parasite transmission to

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native birds via both invasive [novel weapon hypothesis] and migratory species; Marzal et al.

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2018) can be driving the observed patterns. Thus, our results support that House Sparrows can

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act synergistically in relation with urbanization in the species richness decline pattern (Chace and

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Walsh 2006; OrtegaÁlvarez and MacGregor-Fors 2011b, c; Aronson et al. 2014; Sol et al. 2014;

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MacGregor-Fors and García-Arroyo 2017).

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We consider that further directions to test the effects of House Sparrows, in synergy with

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urbanization on native bird communities, require both laboratory and field experiments. In

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doing so, studies ought to consider balanced designs, taking into account diverse urban

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conditions (e.g., residential, industrial, commercial, greenspaces), including non-urban controls

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of different land uses (e.g., original vegetation, agricultural), and several House Sparrow

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abundance scenarios. Additionally, it is of the utmost importance to study House Sparrow

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intraspecific and interspecific interactions (e.g., feeding and nesting resources), as well as

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monitoring their populations in different spatiotemporal scales. Finally, and based on our field

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observations, we highlight the importance of the maintenance of vegetation cover and

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structure in urban areas, not only in large greenspaces but also in private gardens and along

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streets, with the aim of promoting the native avian assemblage diversity.

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Acknowledgments

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We are deeply thankful with María del Coro Arizmendi Arriaga, Roger Guevara, Fabricio

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Villalobos, and José Antonio González Oreja for their helpful comments that enhanced the

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quality and clarity of the manuscript, as well as Miguel Ángel Gómez Martínez, Oscar Humberto

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Marín Gómez, Carlos Mauricio Trujillo Torres, Juan Fernando Escobar Ibáñez, Julian Avila

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Campos, Lorena Ramírez Restrepo, and Sonia Morán for their assistance in the field. MG-A

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acknowledges the scholarship and financial support provided by the National Council of Science

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and Technology (CONACYT 416452) and the Master’s Program of the Instituto de Ecología, A.C.

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(INECOL).

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559
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562

Table 1 Number and distribution of survey sites in the three conditions of the studied urban

563

settlements. a urban House Sparrow invaded, b urban House Sparrow non-invaded, c non-

564

urban House Sparrow non-invaded, d Elevation was retrieved from INEGI (2010)

565
Study region

Settlement

UIa

UNIb

NUNIc

Latitude (N)

Longitude (W)

size (km2)

Elevation (m
a.s.l.)

Xico

567

9

12

21

19° 25′ 21.72”

97° 0′ 33.48”

1320

Teocelo
San Marcos
Úrsulo Galván

566

2
1
0.7
0.14

1
3
0

19
7
4

20
10
4

19° 23′ 7.08”
19° 25′ 22.8”
19° 25′ 45.84”

96° 58′ 30”
96° 57′ 59.04”
96° 58′ 41.88”

1160
1100
1140

568

Table 2 Bird species richness (average ± 84% CI) across seasons in the surveyed conditions

569

considering all studied settlements.

570

a urban House Sparrow invaded, b urban House Sparrow non-invaded, c non-urban House

571

Sparrow non-invaded

572
Season
Condition

Spring

Summer

Fall

Winter

UIa

17.7 ± 4.3

10.7 ± 1.7

11.8 ± 4.4

14.5 ± 6.4

UNIb

19.9 ± 3.6
25.5 ± 3.7

16.0 ± 3.5
21.3 ± 3.5

12.4 ± 2.0
25.6 ± 3.8

16.1 ± 3.2
28.2 ± 3.6

NUNIc

573
574

575
576

Table 3 GAM considering predictor variables describing vegetation characteristics and urban
infrastructure in relation with native bird species richness

577
Variable

579

χ2

P

Season
s (Built cover)
s (Maximum shrub height)
s (House Sparrow abundances)
s (Shrub richness)
s (Passing cars per minute)

578

DF
3
1
1
2
1
1

24.03
28.34
9.13
11.32
0.58
2.54

<0.001
<0.001
0.002
0.012
0.494
0.110

580

Fig. 1 Study areas and sampling locations. Map scales differ for graphical purposes. HS = House

581

Sparrow.

582
583

584

Fig. 2 Bray-Curtis group average link cluster showing avian assemblage composition patterns in

585

the three studied conditions and seasons (UI = urban House Sparrow invaded; UNI = urban

586

House Sparrow non- invaded; NUNI = non-urban House Sparrow non-invaded; numbers after

587

study conditions represent seasons: 1 = spring, 2 = summer, 3 = fall, 4 = winter

588
589
590
591
592

593

Fig. 3 In this graph we display variables that showed to be significantly related with bird species

594

richness in the GAM. Panels a maximum shrub height and b built cover show the relationship

595

with smoothened data, insets represent the best-fit for smoothened and observed values

596

(positive for shrub height, negative for built cover). For c House Sparrow abundances, lower left

597

panel corresponds to the best-fit adjustment, showing the three different scenarios of 0

598

individuals (red), 1–5 individuals (blue), and 6–12 individuals (black). Each scenario connects to

599

its corresponding bird species richness in the lower right panel. Letters below the lower 84% CI

600

bars stand for statistical significant differences. HS = House Sparrow.

601
602

603

Fig. 4 Built cover at the studied conditions. Letters above error bars represent statistical

604

differences

605

606