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The records were gathered from citizen science platforms, field data, and reports from researchers and birdwatchers. A total of 2,618 individuals were recorded across 219 occurrences, 214 of which took place in Rio Grande do Sul, with five others distributed among the states of São Paulo, Rio de Janeiro, and Minas Gerais. In just one decade, the species has already occupied a considerable portion of the Brazilian Pampa, especially in the southernmost regions near the border with Uruguay, with 97.7% of the records concentrated in Rio Grande do Sul. We identified 55 nesting sites, most of which were located in cavities previously excavated by woodpeckers (Picidae), primarily in trees of the genus Eucalyptus . Breeding begins in September, peaks between October and November, and extends until the end of December. The data from this study indicate a high degree of behavioral plasticity in the species, facilitating its rapid adaptation and expansion within the Pampa biome. Sturnidae exotic bird invasive species Pampa biome Figures Figure 1 Figure 2 Figure 3 Introduction Sturnus vulgaris Linnaeus, 1758, commonly called the European starling and a member of the family Sturnidae, is widely recognized as one of the most abundant and impactful invasive bird species globally (Lowe et al. 2000 ). Native to Eurasia and North Africa, it has been introduced to several regions around the world, including the United States, Mexico, South Africa, Australia, New Zealand, Argentina, and various Pacific and Caribbean islands (Craig and Feare 2009 ). In South America, the species was first introduced in Argentina during the 1980s, with nesting records in the Buenos Aires region beginning in 1987 (Pérez 1988 ; Schmidtuz and Agulián 1988). Since then, it has expanded through central and northern Argentina (Peris et al. 2005 ; Zufiaurre et al. 2016 ; Codesido and Drozd 2021 ), reaching Uruguay (Mazzulla 2013 ) and, more recently, Brazil, where it was first recorded in the municipality of Lavras do Sul in the state of Rio Grande do Sul on October 10, 2014 (Silva et al. 2017 ). Invasive species are among the leading causes of biodiversity loss worldwide, and they also generate economic and health-related impacts, including agricultural damage (Vitousek et al. 1996; Wilcove et al. 1998; Lowe et al. 2000 ). The rapid colonization of S. vulgaris in agro-pastoral ecosystems in the Neotropics is facilitated by its dispersal capacity and high ecological plasticity (Peris et al. 2005 ). The species also occupies urban areas, where the diversity of available resources offers favorable conditions for adaptation and expansion (Cadotte et al. 2017 ), potentially serving as hubs for the invasion of adjacent natural habitats (Lockwood et al. 2005 ; Zufiaurre et al. 2016 ). Competition with native birds for nesting sites is one of the main concerns regarding the invasion of S. vulgaris and its effects on local bird communities. Di Giacomo et al. (1993) pointed out that the species nests in tree cavities, light towers, buildings, and artificial nest boxes. During the breeding season, it uses both natural and artificial cavities, such as the nests of Colaptes melanochloros and Colaptes campestris (woodpeckers) and Furnarius rufus (rufous hornero), rooftops, light poles (Rebolo Ifrán and Fiorini 2010; Rizzo 2010 ; Ibáñez 2015). Outside the breeding season, the species gathers in large communal roosts, composed of thousands or even millions of individuals, generally located in urban or semi-urban areas (Feare 1984 ; Clergeau and Quenot 2007 ; Bozzo et al. 2021 ). These roosts are dynamic and seasonal, typically found in dense-canopy trees or urban structures (Symonds 1961; Stewart 1973; Lyon and Caccamise 1981; Clergeau and Quenot 2007 ). In addition to the potential competition for nesting cavities with native birds, the invasion by S. vulgaris may negatively affect agricultural activities and contribute to the transmission of pathogens (Koenig 2003 ; Frei et al. 2015 ; Cabe 2021 ). Therefore, the species represents a significant threat to invaded ecosystems. In this study, we analyzed the expansion of S. vulgaris in Brazil over its first 10 years of colonization, with an emphasis on populations originating from Argentina and Uruguay and present the current status of the species in Brazil. We also aimed to understand its selection of nesting sites, behavioral patterns, and feeding habits as an invasive species in the country. Materials and methods Data collection A comprehensive survey of S. vulgaris records in Brazil was conducted using the main citizen science platforms employed by ornithologists and bird watchers: eBird, iNaturalist, Global Biodiversity Information Facility, and the Brazilian WikiAves. It is important to note that several researchers and birdwatchers, especially users of WikiAves, were directly contacted and provided valuable information through messages on the platform, adding to the records previously published by them. All species records to the end of 2024 were considered. Additionally, field data collected by us, obtained opportunistically during fieldwork and travel along roads, as part of other research projects, mostly ornithological in nature, were included. For each record, the following information was noted: geographic coordinates, number of individuals observed, presence of nesting signs (such as adults visiting nests or chicks observed in the nests), and the observation date. To avoid duplicate records, a filtering process was done for entries present on more than one platform, considering that many records (from different platforms) referred to the same locations and dates. In such cases, only the record with the highest number of individuals per site was counted, thus minimizing the risk of multiple records for certain hotspots, especially in some birdwatching hotspots. We also included information about the species' behavior and feeding habits in Brazil. Statistical analyses The abundance of S. vulgaris over time was analyzed using a Poisson regression model. Data on year of observation and recorded abundance were processed in R (v.4.x.x). The model was fitted with the glm() function using the Poisson family, with abundance as the response variable and year as the predictor. The exponential equation (µ = exp(b0 + b1 * Year)) was derived from the regression coefficients. Model fit was assessed using the Pseudo R² (Deviance), calculated as (null deviance – residual deviance) / null deviance. For visualization, model predictions were generated for a sequence of years, with error bands estimated as the square root of predicted abundance, consistent with the Poisson distribution. The graph, created with ggplot2, displays observed data points, the predicted regression curve, error bands, the model equation, and the Pseudo R² value. For spatial analysis, specialized libraries for visualizing and manipulating geospatial data were used. The ggplot2 package was employed for graph creation, ensuring efficient visualization. The sf (simple features) library was used for the manipulation and analysis of geospatial data in vector format. Data manipulation was performed with dplyr , which provides an intuitive grammar for transformation and analysis. The sp package complemented sf in operations with spatial objects. For geographic data retrieval and creation of interactive maps, ggmap was used, allowing the integration of Google Maps data into the R environment. The creation of the density map involved converting geographic coordinates into a spatial object with sf . Then, the function geom_density_2d() from ggplot2 was applied to represent the density of points, facilitating the identification of areas with higher concentrations and the interpretation of spatial data. RESULTS Species Expansion in Brazil A total of 2,618 individuals were recorded in 219 different occurrences in Brazil, with 214 occurrences in 26 municipalities of Rio Grande do Sul and only five for the rest of the country, located in the states of São Paulo, Rio de Janeiro, and Minas Gerais (Fig. 1 a). The first record of the species in Brazil was made in 2014, in the municipality of Lavras do Sul, Rio Grande do Sul (Silva et al. 2017 ). Since then, in just 10 years, the species has already occupied much of the Brazilian Pampa. The records are mainly concentrated in the extreme south of Brazil, along the border with Uruguay, as the expansion is clearly occurring from south to north, likely with individuals coming from Uruguay and Argentina. Through citizen science platforms, which rely on the collaboration of thousands of birdwatchers and researchers, it was possible to map the progressive distribution of the species in Brazil. The distribution of records reveals a strong concentration in the state of Rio Grande do Sul (97.7%), where spatial expansion has become progressively more evident each year. Based on the compiled data, we generated maps that illustrate this advancement over the last decade (Fig. 1 b). In 2014, there was only one record of the species in the state. After four years, the records had expanded to four municipalities. By 2021, the presence of the species was confirmed in 10 municipalities in Rio Grande do Sul. However, the most concerning growth occurred in the last three years. By 2024, the number of municipalities with records of the species had increased to 26, representing a 150% increase compared to 2021 (Supplementary material). A large portion of these municipalities (24) with records of the species are located in the Brazilian Pampa. However, in recent years, records have also been made in cities within the Atlantic Forest of Rio Grande do Sul, as well as in the southeastern region of Brazil, which likely represent a second source of invasion (independent of the invasion recorded along the southern border). The total area of the 26 municipalities in Rio Grande do Sul with records of S. vulgaris (71,423 km²) represents approximately 36.9% of the total area of the Brazilian Pampa and 25.3% of the area of Rio Grande do Sul (Instituto Brasileiro de Geografia e Estatística 2019). Although the species is not distributed throughout the entire area of the municipalities in which it has been recorded, in several of them, such as Sant’Ana do Livramento, Rio Grande, São Gabriel, and Lavras do Sul, records have been made in two or more locations situated tens of kilometers apart. The Poisson regression analysis revealed a significant positive exponential relationship between the year of observation and the abundance of the exotic species Sturnus vulgaris in Brazil (Fig. 2 ). The regression curve, µ = exp(-1124.16 + 0.56 * Year), indicates accelerated population growth over time. The positive coefficient (0.56) for “Year” suggests an exponential annual increase in abundance. The Pseudo R² of 0.521 shows that the model explains a considerable proportion of the observed variability, indicating a good fit. The widening error bands with increasing abundance are consistent with the Poisson distribution, where variance equals the mean, reflecting greater uncertainty in estimates for larger populations. Despite the overall growth trend, the 2023 data point, with exceptionally high abundance, suggests potential superpopulation events or local variations influencing population dynamics. The 2024 observation, although lower than 2023, remains high, reinforcing the establishment and expansion of the species in Brazil. Nesting Aspects of the Species in Brazil A total of 55 S. vulgaris nesting sites were identified, including sites with multiple nests (Fig. 3 a). The most frequent nesting sites were “tree cavities” (Fig. 3 b) previously created by woodpeckers (Picidae) in trees of the genus Eucalyptus ( n = 43). Additionally, the use of natural cavities in trees such as Alchornea triplinervia , Pinus elliottii , and Melia azedarach was also recorded. In addition to the use of tree cavities and hollows, an interesting behavior was observed on two occasions when the species used the nests of cochicho ( Anumbius annumbi ). In the nests where it was possible to record the number of chicks, the number varied from one to four per nest, with three chicks being apparently the most common During our field observations, an interspecific competition event was recorded on October 3, 2024, when an individual of S. vulgaris was competing for a cavity with a pair of C . campestris that was blocking the entrance to its likely nest in M. azedarach . The species' nesting in the Brazilian Pampa was observed to begin in the austral spring, starting in the second half of September. Our nesting records peaked in October and November, a period that saw the highest occurrence of active nests, extending until the end of December. Confirming these observations, chicks were recorded most frequently in the last three months of the year. Regarding foraging, we observed that flocks of S. vulgaris frequently explored low grasslands in extensive grazing areas, often alongside other species such as Molothrus bonariensis (the shiny cowbird). During parental care, adults captured invertebrates from the ground to feed the juveniles, and they were frequently observed carrying simultaneously in their beaks up to four or five insects, including those from the orders Orthoptera and Lepidoptera. Discussion Sturnus vulgaris is widely recognized as one of the most problematic invasive species in the world. It is currently found on every continent except Antarctica, exhibiting successful dispersal across various regions (Lowe et al. 2000 ). Its expansion rate has been documented in different areas, varying according to environmental and population factors. Since 1987, its average spread in Argentina has been estimated at 7.5 km/year (Peris et al. 2005 ), a relatively low rate when compared to the 43 km/year observed in North America over the first part of the 20th century (Wing 1943 ). More recent observations suggest that this expansion dynamic may be accelerating. In Mar del Plata and Entre Ríos, Argentina, for example, records indicate expansion in a north-south direction at a rate of 26 km/year or 18 km/year, depending on the method of analysis (Peris et al. 2005 ). Models based on European bird banding recovery data suggest that the expansion rate can reach between 46 and 78 km/year (Van den Bosch et al. 1992). Data from Argentina indicate that the species behaves as a partial migrant. During the post-breeding season, the maximum recorded distances from the first sightings in the city of Buenos Aires were 375 km to the south and 400 km to the north, reflecting an estimated winter spread of 25–27 km/year (Peris et al. 2005 ). The invasion by S. vulgaris has proven particularly critical in some regions of southernmost Brazil, especially in the municipalities of Chuí and Santa Vitória do Palmar. Border areas with Uruguay, such as the municipalities of Sant’Ana do Livramento, Bagé, and Aceguá, also concentrate a significant number of records. These localities stand out not only for the high number of occurrences of the species but also for being important nesting areas. This scenario suggests that these regions offer favorable conditions for the reproduction and dispersal of the S. vulgaris , which may facilitate the expansion and establishment of the invasive population. Furthermore, the proximity to Uruguay and Argentina is also an important factor that may facilitate the arrival of new individuals expanding into Brazil in a south–north or west–east direction. Regarding nesting, we recorded S. vulgaris primarily occupying woodpecker cavities, with C. campestris and C. melanochloros being the most common cavity-excavating species in the Brazilian Pampa region, especially in Eucalyptus trees. This behavior has also been reported by other authors, such as Schmidtz and Agulian (1988), who described the occupation of cavities excavated by C. melanochloros in the Argentine provinces of Buenos Aires and Mendoza. Similarly, Peris et al. ( 2005 ), also in Argentina, found that most nests of S. vulgaris were located in cavities made by Colaptes species. In a study conducted in La Plata, Argentina, Palacio et al. ( 2022 ) reported that more than half of the nests (53%) were located in man-made structures, while only 13% were in tree cavities. These findings contrast with those of our study, in which tree cavities were the main nesting sites. It is worth noting, however, that all nests recorded in Brazil were located in rural areas, and there have been no reports of the species nesting in urban environments to date. Palacio et al. ( 2022 ) also found a high frequency of nests built inside the structures of F. rufus (17.3%). In Buenos Aires Province, Rizzo ( 2010 ) also reported the use of F. rufus nests. Although we did not record the use of F. rufus nests in Brazil, it is likely to occur, considering that this is one of the most common bird species in the country. On the other hand, we did observe the use of nests built by A. annumbi , another species of Furnariidae, on two occasions, behavior also recorded by Di Sallo et al. (2014) in Buenos Aires Province. The use of tree cavities for nesting in areas also inhabited by S. vulgaris was observed for several native species, including Drymornis bridgesii , Falco sparverius , and Melanerpes candidus . It is well known that S. vulgaris competes for nesting sites with native species in areas where it has been introduced (Chow 2000; Ruda Veja 2004 ; Gómez de Silva 2005; Peris et al. 2005 ; Rizzo 2010 ). In Australia and Europe, studies have reported competition for nesting sites (Pazzucconi 1997; Pell and Tidemann 1997; Wiebe 2003). It is likely that some native cavity-nesting species may face challenges in securing nesting sites due to competition with S. vulgaris , such as Sicalis flaveola and Progne tapera (Rizzo 2010 ). The nests of S. vulgaris were found in four different tree taxa. Preference for a specific tree species was observed in La Plata, where all trees used by S. vulgaris belonged to evergreen species, particularly the palm Phoenix canariensis , which is known for its dense foliage and high canopy. Girini et al. ( 2014 ) reported the presence of nests in various tree species, including those from the genera Ulmus , Populus , Eucalyptus , and Quercus . Records of nesting in natural environments span several regions of Argentina, including Buenos Aires, Córdoba, and Mendoza (Schmidtz and Agulian 1988; Klavins and Álvarez 2012 ; Zanotti 2013 ). On the other hand, Di Giacomo et al. (1993) mentioned that nesting sites may include not only tree cavities but also lighting towers, buildings, wall cavities, and artificial nest boxes. In Brazil, the use of these substrates by S. vulgaris is not yet recorded, possibly because the birds have primarily been observed in rural and pastoral areas and have not, so far, established populations in urban centers. This scenario contrasts with observations in Argentina, where, for example, roosts with more than 1,000 individuals were reported in the city of La Plata (Girini et al. 2014 ). The reproductive period observed in Brazil is similar to that in Argentina, where the species also begins breeding activity in September, extending through December (Rebolo Ifran and Fiorini 2010 ), with nestlings observed in October (Schmidtz and Agulian 1988) and juveniles recorded between November and January (Peris et al. 2005 ; Rizzo 2010 ). We observed parents feeding chicks mainly with invertebrates from the orders Orthoptera and Lepidoptera. After fledging, juveniles may shift to a predominantly frugivorous diet, forming large flocks that attack a variety of crops, including grapes, strawberries, apples, cherries, blueberries, and other small fruits (Conover and Dolbeer 2007 ; Linz et al. 2007 ). The invasion of S. vulgaris likely poses a direct threat to grape and olive production in southern Brazil, as large flocks can become a significant pest for soft fruit crops (Weber 1979 ; Chow 2000; Gómez de Silva 2005). In North America, blueberry crops are among the most affected by S. vulgaris , along with grapes, olives, and cherries (Knittle and Guarino 1976 ). Pimentel (2000) estimated that the damage caused by this species to U.S. agriculture reaches approximately US $ 800 million per year, highlighting the seriousness of the problem. This figure includes not only direct production losses but also additional costs associated with control and mitigation strategies, such as protective netting, acoustic deterrents, and population management programs. Given the species’ documented ecological and agricultural effects, it is essential to deepen our understanding of the ecology of S. vulgaris in the Neotropical region. Future studies should more thoroughly investigate its effects on native species, as well as its reproductive and feeding biology, allowing for a clearer understanding of its impacts on the natural environments it occupies. Furthermore, considering its ongoing geographic and population expansion, continued research will be crucial to monitor its population dynamics and expansion patterns in native habitats. The potential impact of this species already suggests the need to implement control programs, particularly in the Brazilian Pampa and southern Brazil, which currently represent the main frontier of the species’ expansion. Declarations Funding declaration Funding for this project was provided by doctoral scholarships granted by the “Coordenação de Aperfeiçoamento de Pessoal de Nível Superior –Brazil (CAPES) – under Funding Code 001." Author Contribution All authors contributed to the conception and design of the study, as well as to field records of the focal species. Metadata collection and organization were performed by José Paulo Souto Dias and Luciano Lopes Marques. Statistical analyses were performed by Jorge Renato Pinheiro Velloso and José Paulo Souto Dias. The first draft of the manuscript was written by José Paulo Souto Dias, Jorge Renato Pinheiro Velloso e Fernanda Machado Teixeira e revised and edited by Luciano Lopes Marques, Cassiana Alves de Aguiar and Carlos Benhur Kasper. All authors read and approved the final manuscript, the author order, and the designation of the corresponding author. Acknowledgement We are deeply grateful to all researchers, biologists, and birdwatchers who shared information about their records of the species, making this study possible. We also thank our colleagues from the GOA Macá birdwatching group who, even from a distance, contributed valuable data. This work was supported by the "Coordination for the Improvement of Higher Education Personnel" (CAPES), Brazil, Funding Code 001. References Aphalo PJ (2020) ggpmisc: Miscellaneous Functions for ‘ggplot2’. R package version 0.3.7, 2020. https://cran.rproject.org/web/packages/ggpmisc/index.html. 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The Auk 60:74-87 Zanotti M (2013) Presencia del Estornino Pinto ( Sturnus vulgaris ) en la provincia de Mendoza, Argentina. Nuestras Aves 58:5–7 Zufiaurre E, Abba A, Bilenca D, Codesido M (2016) Role of landscape elements on recent distributional expansion of European starlings ( Sturnus vulgaris ) in agroecosystems of the Pampas, Argentina. Wilson J of Ornithol 128:306–313. https://doi.org/10.1676/wils-128-02-306-313.1 Additional Declarations No competing interests reported. Supplementary Files SupplementarymaterialSturnus.tiff Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Dias","email":"data:image/png;base64,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","orcid":"","institution":"Universidade Federal do Pampa","correspondingAuthor":true,"prefix":"","firstName":"José","middleName":"Paulo Souto","lastName":"Dias","suffix":""},{"id":539858804,"identity":"c01ed615-3180-471f-b09c-4a9a1d19e020","order_by":1,"name":"Jorge Renato Pinheiro Velloso","email":"","orcid":"","institution":"Universidade Federal do Pampa","correspondingAuthor":false,"prefix":"","firstName":"Jorge","middleName":"Renato Pinheiro","lastName":"Velloso","suffix":""},{"id":539858805,"identity":"051b5666-65b6-423c-b557-81f53022b1ca","order_by":2,"name":"Fernanda Machado-Teixeira","email":"","orcid":"","institution":"Universidade Federal do Pampa","correspondingAuthor":false,"prefix":"","firstName":"Fernanda","middleName":"","lastName":"Machado-Teixeira","suffix":""},{"id":539858806,"identity":"48d3a84e-5fdf-44ce-a8f3-93419fe666fe","order_by":3,"name":"Luciano Lopes Marques","email":"","orcid":"","institution":"Universidade Federal do Pampa","correspondingAuthor":false,"prefix":"","firstName":"Luciano","middleName":"Lopes","lastName":"Marques","suffix":""},{"id":539858807,"identity":"3b2ecb73-4a73-48de-86e1-782e5b635502","order_by":4,"name":"Cassiana Alves de Aguiar","email":"","orcid":"","institution":"Instituto de Conservação Eco dos Campos","correspondingAuthor":false,"prefix":"","firstName":"Cassiana","middleName":"Alves","lastName":"de Aguiar","suffix":""},{"id":539858808,"identity":"c03a7ef7-50a1-4813-94f5-15391b119308","order_by":5,"name":"Carlos Benhur Kasper","email":"","orcid":"","institution":"Universidade Federal do Pampa","correspondingAuthor":false,"prefix":"","firstName":"Carlos","middleName":"Benhur","lastName":"Kasper","suffix":""}],"badges":[],"createdAt":"2025-09-09 13:23:43","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7574299/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7574299/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":95313253,"identity":"eed71ccf-0b46-4ea6-b2e5-eeee6d90a143","added_by":"auto","created_at":"2025-11-06 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07:16:15","extension":"html","order_by":20,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":97891,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7574299/v1/8bdc3306d024b564265d64c0.html"},{"id":95313366,"identity":"050aa0c5-1c4d-4e15-8ebd-b5d37adb9451","added_by":"auto","created_at":"2025-11-06 15:51:18","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1537945,"visible":true,"origin":"","legend":"\u003cp\u003e(a) Current distribution of \u003cem\u003eSturnus vulgaris\u003c/em\u003e in Brazil. (b) Spatial expansion of the species in Rio Grande do Sul from 2014 to 2024. The size and color of the dots represent the number of recorded individuals, with a gradient from light green (fewer individuals) to dark green/black (more individuals).\u003c/p\u003e","description":"","filename":"Fig1.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7574299/v1/fd2b418a7be27b898f865bed.jpg"},{"id":95271113,"identity":"e1d0fb06-8780-4738-99cf-2a560887ba0b","added_by":"auto","created_at":"2025-11-06 07:16:15","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":806339,"visible":true,"origin":"","legend":"\u003cp\u003ePoisson regression of \u003cem\u003eSturnus vulgaris\u003c/em\u003e (European starling) abundance in Brazil as a function of observation year. Black points represent observed abundance data. The solid blue line indicates the abundance curve predicted by the Poisson regression model, with the equation μ = exp(-1124.16 + 0.56 * Year) and a Pseudo R² of 0.521. The gray shaded area represents the error band (±1 standard deviation) of the prediction, reflecting the uncertainty associated with the abundance estimate. The X-axis represents the year of observation and the Y-axis represents the abundance of individuals.\u003c/p\u003e","description":"","filename":"Fig2.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7574299/v1/a3c64f3410ff7c45874bad74.jpg"},{"id":95271115,"identity":"4d7b6180-0e54-41f9-aeb2-171bc83f36b7","added_by":"auto","created_at":"2025-11-06 07:16:15","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1691017,"visible":true,"origin":"","legend":"\u003cp\u003e(a) Nesting record locations of \u003cem\u003eSturnus vulgaris\u003c/em\u003e in the state of Rio Grande do Sul, southern Brazil. (b) Adult of \u003cem\u003eS. vulgaris\u003c/em\u003e at the entrance of a nest, recorded in the state of Rio Grande do Sul. Photo credit: J. P. S. Dias.\u003c/p\u003e","description":"","filename":"fig3.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7574299/v1/c4ed3363eead333e60da57e6.jpg"},{"id":105566333,"identity":"5b9009f1-ce12-4a59-bd23-11917d2f33df","added_by":"auto","created_at":"2026-03-27 12:56:11","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4537790,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7574299/v1/17523133-935e-4380-bf63-c39eb3874623.pdf"},{"id":95271118,"identity":"29bfc0a6-6e5b-426f-93ea-7355ba7a6602","added_by":"auto","created_at":"2025-11-06 07:16:15","extension":"tiff","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":2895138,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementarymaterialSturnus.tiff","url":"https://assets-eu.researchsquare.com/files/rs-7574299/v1/e6df638c9991059a03001ea8.tiff"}],"financialInterests":"No competing interests reported.","formattedTitle":"Ten years of the European Starling (Sturnus vulgaris) in Brazil: range expansion and reproductive insights","fulltext":[{"header":"Introduction","content":"\u003cp\u003e\u003cem\u003eSturnus vulgaris\u003c/em\u003e Linnaeus, 1758, commonly called the European starling and a member of the family Sturnidae, is widely recognized as one of the most abundant and impactful invasive bird species globally (Lowe et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Native to Eurasia and North Africa, it has been introduced to several regions around the world, including the United States, Mexico, South Africa, Australia, New Zealand, Argentina, and various Pacific and Caribbean islands (Craig and Feare \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). In South America, the species was first introduced in Argentina during the 1980s, with nesting records in the Buenos Aires region beginning in 1987 (P\u0026eacute;rez \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Schmidtuz and Aguli\u0026aacute;n 1988). Since then, it has expanded through central and northern Argentina (Peris et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Zufiaurre et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Codesido and Drozd \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), reaching Uruguay (Mazzulla \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and, more recently, Brazil, where it was first recorded in the municipality of Lavras do Sul in the state of Rio Grande do Sul on October 10, 2014 (Silva et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eInvasive species are among the leading causes of biodiversity loss worldwide, and they also generate economic and health-related impacts, including agricultural damage (Vitousek et al. 1996; Wilcove et al. 1998; Lowe et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). The rapid colonization of \u003cem\u003eS. vulgaris\u003c/em\u003e in agro-pastoral ecosystems in the Neotropics is facilitated by its dispersal capacity and high ecological plasticity (Peris et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). The species also occupies urban areas, where the diversity of available resources offers favorable conditions for adaptation and expansion (Cadotte et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), potentially serving as hubs for the invasion of adjacent natural habitats (Lockwood et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Zufiaurre et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eCompetition with native birds for nesting sites is one of the main concerns regarding the invasion of \u003cem\u003eS. vulgaris\u003c/em\u003e and its effects on local bird communities. Di Giacomo et al. (1993) pointed out that the species nests in tree cavities, light towers, buildings, and artificial nest boxes. During the breeding season, it uses both natural and artificial cavities, such as the nests of \u003cem\u003eColaptes melanochloros\u003c/em\u003e and \u003cem\u003eColaptes campestris\u003c/em\u003e (woodpeckers) and \u003cem\u003eFurnarius rufus\u003c/em\u003e (rufous hornero), rooftops, light poles (Rebolo Ifr\u0026aacute;n and Fiorini 2010; Rizzo \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Ib\u0026aacute;\u0026ntilde;ez 2015). Outside the breeding season, the species gathers in large communal roosts, composed of thousands or even millions of individuals, generally located in urban or semi-urban areas (Feare \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Clergeau and Quenot \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Bozzo et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These roosts are dynamic and seasonal, typically found in dense-canopy trees or urban structures (Symonds 1961; Stewart 1973; Lyon and Caccamise 1981; Clergeau and Quenot \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn addition to the potential competition for nesting cavities with native birds, the invasion by \u003cem\u003eS. vulgaris\u003c/em\u003e may negatively affect agricultural activities and contribute to the transmission of pathogens (Koenig \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Frei et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Cabe \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Therefore, the species represents a significant threat to invaded ecosystems. In this study, we analyzed the expansion of \u003cem\u003eS. vulgaris\u003c/em\u003e in Brazil over its first 10 years of colonization, with an emphasis on populations originating from Argentina and Uruguay and present the current status of the species in Brazil. We also aimed to understand its selection of nesting sites, behavioral patterns, and feeding habits as an invasive species in the country.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eData collection\u003c/h2\u003e\u003cp\u003eA comprehensive survey of \u003cem\u003eS. vulgaris\u003c/em\u003e records in Brazil was conducted using the main citizen science platforms employed by ornithologists and bird watchers: eBird, iNaturalist, Global Biodiversity Information Facility, and the Brazilian WikiAves. It is important to note that several researchers and birdwatchers, especially users of WikiAves, were directly contacted and provided valuable information through messages on the platform, adding to the records previously published by them. All species records to the end of 2024 were considered. Additionally, field data collected by us, obtained opportunistically during fieldwork and travel along roads, as part of other research projects, mostly ornithological in nature, were included.\u003c/p\u003e\u003cp\u003eFor each record, the following information was noted: geographic coordinates, number of individuals observed, presence of nesting signs (such as adults visiting nests or chicks observed in the nests), and the observation date. To avoid duplicate records, a filtering process was done for entries present on more than one platform, considering that many records (from different platforms) referred to the same locations and dates. In such cases, only the record with the highest number of individuals per site was counted, thus minimizing the risk of multiple records for certain hotspots, especially in some birdwatching hotspots. We also included information about the species' behavior and feeding habits in Brazil.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eStatistical analyses\u003c/h3\u003e\n\u003cp\u003eThe abundance of \u003cem\u003eS. vulgaris\u003c/em\u003e over time was analyzed using a Poisson regression model. Data on year of observation and recorded abundance were processed in R (v.4.x.x). The model was fitted with the glm() function using the Poisson family, with abundance as the response variable and year as the predictor. The exponential equation (\u0026micro;\u0026thinsp;=\u0026thinsp;exp(b0\u0026thinsp;+\u0026thinsp;b1 * Year)) was derived from the regression coefficients. Model fit was assessed using the Pseudo R\u0026sup2; (Deviance), calculated as (null deviance \u0026ndash; residual deviance) / null deviance. For visualization, model predictions were generated for a sequence of years, with error bands estimated as the square root of predicted abundance, consistent with the Poisson distribution. The graph, created with ggplot2, displays observed data points, the predicted regression curve, error bands, the model equation, and the Pseudo R\u0026sup2; value.\u003c/p\u003e\u003cp\u003eFor spatial analysis, specialized libraries for visualizing and manipulating geospatial data were used. The \u003cem\u003eggplot2\u003c/em\u003e package was employed for graph creation, ensuring efficient visualization. The \u003cem\u003esf\u003c/em\u003e (simple features) library was used for the manipulation and analysis of geospatial data in vector format. Data manipulation was performed with \u003cem\u003edplyr\u003c/em\u003e, which provides an intuitive grammar for transformation and analysis. The \u003cem\u003esp\u003c/em\u003e package complemented \u003cem\u003esf\u003c/em\u003e in operations with spatial objects. For geographic data retrieval and creation of interactive maps, \u003cem\u003eggmap\u003c/em\u003e was used, allowing the integration of Google Maps data into the R environment.\u003c/p\u003e\u003cp\u003eThe creation of the density map involved converting geographic coordinates into a spatial object with \u003cem\u003esf\u003c/em\u003e. Then, the function \u003cem\u003egeom_density_2d()\u003c/em\u003e from \u003cem\u003eggplot2\u003c/em\u003e was applied to represent the density of points, facilitating the identification of areas with higher concentrations and the interpretation of spatial data.\u003c/p\u003e"},{"header":"RESULTS","content":"\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e\u003ch2\u003eSpecies Expansion in Brazil\u003c/h2\u003e\u003cp\u003eA total of 2,618 individuals were recorded in 219 different occurrences in Brazil, with 214 occurrences in 26 municipalities of Rio Grande do Sul and only five for the rest of the country, located in the states of S\u0026atilde;o Paulo, Rio de Janeiro, and Minas Gerais (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea). The first record of the species in Brazil was made in 2014, in the municipality of Lavras do Sul, Rio Grande do Sul (Silva et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Since then, in just 10 years, the species has already occupied much of the Brazilian Pampa. The records are mainly concentrated in the extreme south of Brazil, along the border with Uruguay, as the expansion is clearly occurring from south to north, likely with individuals coming from Uruguay and Argentina.\u003c/p\u003e\u003cp\u003eThrough citizen science platforms, which rely on the collaboration of thousands of birdwatchers and researchers, it was possible to map the progressive distribution of the species in Brazil. The distribution of records reveals a strong concentration in the state of Rio Grande do Sul (97.7%), where spatial expansion has become progressively more evident each year. Based on the compiled data, we generated maps that illustrate this advancement over the last decade (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb).\u003c/p\u003e\u003cp\u003eIn 2014, there was only one record of the species in the state. After four years, the records had expanded to four municipalities. By 2021, the presence of the species was confirmed in 10 municipalities in Rio Grande do Sul. However, the most concerning growth occurred in the last three years. By 2024, the number of municipalities with records of the species had increased to 26, representing a 150% increase compared to 2021 (Supplementary material). A large portion of these municipalities (24) with records of the species are located in the Brazilian Pampa. However, in recent years, records have also been made in cities within the Atlantic Forest of Rio Grande do Sul, as well as in the southeastern region of Brazil, which likely represent a second source of invasion (independent of the invasion recorded along the southern border).\u003c/p\u003e\u003cp\u003eThe total area of the 26 municipalities in Rio Grande do Sul with records of \u003cem\u003eS. vulgaris\u003c/em\u003e (71,423 km\u0026sup2;) represents approximately 36.9% of the total area of the Brazilian Pampa and 25.3% of the area of Rio Grande do Sul (Instituto Brasileiro de Geografia e Estat\u0026iacute;stica 2019). Although the species is not distributed throughout the entire area of the municipalities in which it has been recorded, in several of them, such as Sant\u0026rsquo;Ana do Livramento, Rio Grande, S\u0026atilde;o Gabriel, and Lavras do Sul, records have been made in two or more locations situated tens of kilometers apart.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eThe Poisson regression analysis revealed a significant positive exponential relationship between the year of observation and the abundance of the exotic species \u003cem\u003eSturnus vulgaris\u003c/em\u003e in Brazil (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The regression curve, \u0026micro;\u0026thinsp;=\u0026thinsp;exp(-1124.16\u0026thinsp;+\u0026thinsp;0.56 * Year), indicates accelerated population growth over time. The positive coefficient (0.56) for \u0026ldquo;Year\u0026rdquo; suggests an exponential annual increase in abundance. The Pseudo R\u0026sup2; of 0.521 shows that the model explains a considerable proportion of the observed variability, indicating a good fit. The widening error bands with increasing abundance are consistent with the Poisson distribution, where variance equals the mean, reflecting greater uncertainty in estimates for larger populations. Despite the overall growth trend, the 2023 data point, with exceptionally high abundance, suggests potential superpopulation events or local variations influencing population dynamics. The 2024 observation, although lower than 2023, remains high, reinforcing the establishment and expansion of the species in Brazil.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eNesting Aspects of the Species in Brazil\u003c/h3\u003e\n\u003cp\u003eA total of 55 \u003cem\u003eS. vulgaris\u003c/em\u003e nesting sites were identified, including sites with multiple nests (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea). The most frequent nesting sites were \u0026ldquo;tree cavities\u0026rdquo; (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb) previously created by woodpeckers (Picidae) in trees of the genus \u003cem\u003eEucalyptus\u003c/em\u003e (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;43). Additionally, the use of natural cavities in trees such as \u003cem\u003eAlchornea triplinervia\u003c/em\u003e, \u003cem\u003ePinus elliottii\u003c/em\u003e, and \u003cem\u003eMelia azedarach\u003c/em\u003e was also recorded. In addition to the use of tree cavities and hollows, an interesting behavior was observed on two occasions when the species used the nests of \u003cem\u003ecochicho\u003c/em\u003e (\u003cem\u003eAnumbius annumbi\u003c/em\u003e). In the nests where it was possible to record the number of chicks, the number varied from one to four per nest, with three chicks being apparently the most common During our field observations, an interspecific competition event was recorded on October 3, 2024, when an individual of \u003cem\u003eS. vulgaris\u003c/em\u003e was competing for a cavity with a pair of \u003cem\u003eC\u003c/em\u003e. \u003cem\u003ecampestris\u003c/em\u003e that was blocking the entrance to its likely nest in \u003cem\u003eM. azedarach\u003c/em\u003e.\u003c/p\u003e\u003cp\u003eThe species' nesting in the Brazilian Pampa was observed to begin in the austral spring, starting in the second half of September. Our nesting records peaked in October and November, a period that saw the highest occurrence of active nests, extending until the end of December. Confirming these observations, chicks were recorded most frequently in the last three months of the year.\u003c/p\u003e\u003cp\u003eRegarding foraging, we observed that flocks of \u003cem\u003eS. vulgaris\u003c/em\u003e frequently explored low grasslands in extensive grazing areas, often alongside other species such as \u003cem\u003eMolothrus bonariensis\u003c/em\u003e (the shiny cowbird). During parental care, adults captured invertebrates from the ground to feed the juveniles, and they were frequently observed carrying simultaneously in their beaks up to four or five insects, including those from the orders Orthoptera and Lepidoptera.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003e\u003cem\u003eSturnus vulgaris\u003c/em\u003e is widely recognized as one of the most problematic invasive species in the world. It is currently found on every continent except Antarctica, exhibiting successful dispersal across various regions (Lowe et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Its expansion rate has been documented in different areas, varying according to environmental and population factors. Since 1987, its average spread in Argentina has been estimated at 7.5 km/year (Peris et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), a relatively low rate when compared to the 43 km/year observed in North America over the first part of the 20th century (Wing \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e1943\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eMore recent observations suggest that this expansion dynamic may be accelerating. In Mar del Plata and Entre R\u0026iacute;os, Argentina, for example, records indicate expansion in a north-south direction at a rate of 26 km/year or 18 km/year, depending on the method of analysis (Peris et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Models based on European bird banding recovery data suggest that the expansion rate can reach between 46 and 78 km/year (Van den Bosch et al. 1992). Data from Argentina indicate that the species behaves as a partial migrant. During the post-breeding season, the maximum recorded distances from the first sightings in the city of Buenos Aires were 375 km to the south and 400 km to the north, reflecting an estimated winter spread of 25\u0026ndash;27 km/year (Peris et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2005\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe invasion by \u003cem\u003eS. vulgaris\u003c/em\u003e has proven particularly critical in some regions of southernmost Brazil, especially in the municipalities of Chu\u0026iacute; and Santa Vit\u0026oacute;ria do Palmar. Border areas with Uruguay, such as the municipalities of Sant\u0026rsquo;Ana do Livramento, Bag\u0026eacute;, and Acegu\u0026aacute;, also concentrate a significant number of records. These localities stand out not only for the high number of occurrences of the species but also for being important nesting areas. This scenario suggests that these regions offer favorable conditions for the reproduction and dispersal of the \u003cem\u003eS. vulgaris\u003c/em\u003e, which may facilitate the expansion and establishment of the invasive population. Furthermore, the proximity to Uruguay and Argentina is also an important factor that may facilitate the arrival of new individuals expanding into Brazil in a south\u0026ndash;north or west\u0026ndash;east direction.\u003c/p\u003e\u003cp\u003eRegarding nesting, we recorded \u003cem\u003eS. vulgaris\u003c/em\u003e primarily occupying woodpecker cavities, with \u003cem\u003eC. campestris\u003c/em\u003e and \u003cem\u003eC. melanochloros\u003c/em\u003e being the most common cavity-excavating species in the Brazilian Pampa region, especially in \u003cem\u003eEucalyptus\u003c/em\u003e trees. This behavior has also been reported by other authors, such as Schmidtz and Agulian (1988), who described the occupation of cavities excavated by \u003cem\u003eC. melanochloros\u003c/em\u003e in the Argentine provinces of Buenos Aires and Mendoza. Similarly, Peris et al. (\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), also in Argentina, found that most nests of \u003cem\u003eS. vulgaris\u003c/em\u003e were located in cavities made by \u003cem\u003eColaptes\u003c/em\u003e species.\u003c/p\u003e\u003cp\u003eIn a study conducted in La Plata, Argentina, Palacio et al. (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) reported that more than half of the nests (53%) were located in man-made structures, while only 13% were in tree cavities. These findings contrast with those of our study, in which tree cavities were the main nesting sites. It is worth noting, however, that all nests recorded in Brazil were located in rural areas, and there have been no reports of the species nesting in urban environments to date. Palacio et al. (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) also found a high frequency of nests built inside the structures of \u003cem\u003eF. rufus\u003c/em\u003e (17.3%). In Buenos Aires Province, Rizzo (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) also reported the use of \u003cem\u003eF. rufus\u003c/em\u003e nests. Although we did not record the use of \u003cem\u003eF. rufus\u003c/em\u003e nests in Brazil, it is likely to occur, considering that this is one of the most common bird species in the country. On the other hand, we did observe the use of nests built by \u003cem\u003eA. annumbi\u003c/em\u003e, another species of Furnariidae, on two occasions, behavior also recorded by Di Sallo et al. (2014) in Buenos Aires Province.\u003c/p\u003e\u003cp\u003eThe use of tree cavities for nesting in areas also inhabited by \u003cem\u003eS. vulgaris\u003c/em\u003e was observed for several native species, including \u003cem\u003eDrymornis bridgesii\u003c/em\u003e, \u003cem\u003eFalco sparverius\u003c/em\u003e, and \u003cem\u003eMelanerpes candidus\u003c/em\u003e. It is well known that \u003cem\u003eS. vulgaris\u003c/em\u003e competes for nesting sites with native species in areas where it has been introduced (Chow 2000; Ruda Veja \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; G\u0026oacute;mez de Silva 2005; Peris et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Rizzo \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). In Australia and Europe, studies have reported competition for nesting sites (Pazzucconi 1997; Pell and Tidemann 1997; Wiebe 2003). It is likely that some native cavity-nesting species may face challenges in securing nesting sites due to competition with \u003cem\u003eS. vulgaris\u003c/em\u003e, such as \u003cem\u003eSicalis flaveola\u003c/em\u003e and \u003cem\u003eProgne tapera\u003c/em\u003e (Rizzo \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2010\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe nests of \u003cem\u003eS. vulgaris\u003c/em\u003e were found in four different tree taxa. Preference for a specific tree species was observed in La Plata, where all trees used by \u003cem\u003eS. vulgaris\u003c/em\u003e belonged to evergreen species, particularly the palm \u003cem\u003ePhoenix canariensis\u003c/em\u003e, which is known for its dense foliage and high canopy. Girini et al. (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) reported the presence of nests in various tree species, including those from the genera \u003cem\u003eUlmus\u003c/em\u003e, \u003cem\u003ePopulus\u003c/em\u003e, \u003cem\u003eEucalyptus\u003c/em\u003e, and \u003cem\u003eQuercus\u003c/em\u003e. Records of nesting in natural environments span several regions of Argentina, including Buenos Aires, C\u0026oacute;rdoba, and Mendoza (Schmidtz and Agulian 1988; Klavins and \u0026Aacute;lvarez \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Zanotti \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). On the other hand, Di Giacomo et al. (1993) mentioned that nesting sites may include not only tree cavities but also lighting towers, buildings, wall cavities, and artificial nest boxes. In Brazil, the use of these substrates by \u003cem\u003eS. vulgaris\u003c/em\u003e is not yet recorded, possibly because the birds have primarily been observed in rural and pastoral areas and have not, so far, established populations in urban centers. This scenario contrasts with observations in Argentina, where, for example, roosts with more than 1,000 individuals were reported in the city of La Plata (Girini et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe reproductive period observed in Brazil is similar to that in Argentina, where the species also begins breeding activity in September, extending through December (Rebolo Ifran and Fiorini \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), with nestlings observed in October (Schmidtz and Agulian 1988) and juveniles recorded between November and January (Peris et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Rizzo \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). We observed parents feeding chicks mainly with invertebrates from the orders Orthoptera and Lepidoptera. After fledging, juveniles may shift to a predominantly frugivorous diet, forming large flocks that attack a variety of crops, including grapes, strawberries, apples, cherries, blueberries, and other small fruits (Conover and Dolbeer \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Linz et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The invasion of \u003cem\u003eS. vulgaris\u003c/em\u003e likely poses a direct threat to grape and olive production in southern Brazil, as large flocks can become a significant pest for soft fruit crops (Weber \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1979\u003c/span\u003e; Chow 2000; G\u0026oacute;mez de Silva 2005). In North America, blueberry crops are among the most affected by \u003cem\u003eS. vulgaris\u003c/em\u003e, along with grapes, olives, and cherries (Knittle and Guarino \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1976\u003c/span\u003e). Pimentel (2000) estimated that the damage caused by this species to U.S. agriculture reaches approximately US\u003cspan\u003e$\u003c/span\u003e800\u0026nbsp;million per year, highlighting the seriousness of the problem. This figure includes not only direct production losses but also additional costs associated with control and mitigation strategies, such as protective netting, acoustic deterrents, and population management programs.\u003c/p\u003e\u003cp\u003eGiven the species\u0026rsquo; documented ecological and agricultural effects, it is essential to deepen our understanding of the ecology of \u003cem\u003eS. vulgaris\u003c/em\u003e in the Neotropical region. Future studies should more thoroughly investigate its effects on native species, as well as its reproductive and feeding biology, allowing for a clearer understanding of its impacts on the natural environments it occupies. Furthermore, considering its ongoing geographic and population expansion, continued research will be crucial to monitor its population dynamics and expansion patterns in native habitats. The potential impact of this species already suggests the need to implement control programs, particularly in the Brazilian Pampa and southern Brazil, which currently represent the main frontier of the species\u0026rsquo; expansion.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eFunding declaration\u003c/h2\u003e\u003cp\u003eFunding for this project was provided by doctoral scholarships granted by the \u0026ldquo;Coordena\u0026ccedil;\u0026atilde;o de Aperfei\u0026ccedil;oamento de Pessoal de N\u0026iacute;vel Superior \u0026ndash;Brazil (CAPES) \u0026ndash; under Funding Code 001.\"\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eAll authors contributed to the conception and design of the study, as well as to field records of the focal species. Metadata collection and organization were performed by Jos\u0026eacute; Paulo Souto Dias and Luciano Lopes Marques. Statistical analyses were performed by Jorge Renato Pinheiro Velloso and Jos\u0026eacute; Paulo Souto Dias. The first draft of the manuscript was written by Jos\u0026eacute; Paulo Souto Dias, Jorge Renato Pinheiro Velloso e Fernanda Machado Teixeira e revised and edited by Luciano Lopes Marques, Cassiana Alves de Aguiar and Carlos Benhur Kasper. All authors read and approved the final manuscript, the author order, and the designation of the corresponding author.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe are deeply grateful to all researchers, biologists, and birdwatchers who shared information about their records of the species, making this study possible. We also thank our colleagues from the GOA Mac\u0026aacute; birdwatching group who, even from a distance, contributed valuable data. This work was supported by the \"Coordination for the Improvement of Higher Education Personnel\" (CAPES), Brazil, Funding Code 001.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAphalo PJ (2020) ggpmisc: Miscellaneous Functions for \u0026lsquo;ggplot2\u0026rsquo;. R package version 0.3.7, 2020. https://cran.rproject.org/web/packages/ggpmisc/index.html. Accessed 14 Jan 2025\u003c/li\u003e\n\u003cli\u003eAzpiroz AB (2012) Aves de las pampas y campos de Argentina, Brasil y Uruguai: una gu\u0026iacute;a de identificaci\u0026oacute;n. Pressur, Nueva Helvecia\u003c/li\u003e\n\u003cli\u003eBivand RS, Gomez-Rubio V (2019) Spatial Data Analysis with R. Springer. \u003c/li\u003e\n\u003cli\u003eBozzo F, Tarricone S, Petrontino A, Cagnetta P, Maringelli L, La Gioia F, Fuccii V, Ragni M (2021) Quantification of the Starling population, estimation and mapping of the damage to olive crops in the Apulia region. Anim 11:1119. https://doi.org/10.3390/ani11041119\u003c/li\u003e\n\u003cli\u003eCabe PR (2021) European starlings (\u003cem\u003eSturnus vulgaris\u003c/em\u003e) as vectors and reservoirs of pathogens affecting humans and domestic livestock. Anim 11:4. https://doi.org/10.3390/ani11020466\u003c/li\u003e\n\u003cli\u003eCadotte MW, Yasui SLE, Livingstone S, Macivor JS\u003cem\u003e \u003c/em\u003e(2017) Are urban systems beneficial, detrimental, or indifferent for biological invasion? Biol Invasions 19:3489\u0026ndash;3503. https://doi.org/10.1007/s10530-017-1586-y Chow J (2000) \u003cem\u003eSturnus vulgaris\u003c/em\u003e. Animal Diversity Web. http://animaldiversity.ummz.umich.edu/accounts/Sturnus_vulgaris/. Accessed 14 Jan 2025\u003c/li\u003e\n\u003cli\u003eClergeau P, Quenot F (2007) Roost selection flexibility of European starlings aids invasion of urban landscape. Landsc and Urban Plan 80:56. https://doi.org/10.1016/j.landurbplan.2006.06.002\u003c/li\u003e\n\u003cli\u003eCodesido M, Drozd A (2021) Alien birds in Argentina: pathways, characteristics and ecological roles. Biol Invasions 23:1329\u0026ndash;1338. https://doi.org/10.1007/s10530-020-02444-w \u003c/li\u003e\n\u003cli\u003eConover MR, Dolbeer RA (2007) Use of decoy traps to protect blueberries from juvenile European starlings. Hum-Wildl Confl 2:265\u0026ndash;270\u003c/li\u003e\n\u003cli\u003eCraig AJ, Feare CJ (2009) Family Sturnidae (starlings). In: del Hoyo J, Elliot A, Christie DA (eds) Handbook of the birds of the world Volume 14. Lynx Edicions, Barcelona, pp 654\u0026ndash;758\u003c/li\u003e\n\u003cli\u003eDe La Pe\u0026ntilde;a MR, Rumboll M (1998) Birds of southern South America and Antarctica Volume 28. Princeton University Press, New Jersey\u003c/li\u003e\n\u003cli\u003eDi Giacoo G, Giacoo I, Abasas M (1993) Nuevos registros de \u003cem\u003eSturnus vulgaris\u003c/em\u003e y \u003cem\u003eAcridotheres cristatellus\u003c/em\u003e en Buenos Aires. Nuestras Aves 29:32\u0026ndash;33\u003c/li\u003e\n\u003cli\u003eDi Sallo FG, Segura LN (2014) Nidificaci\u0026oacute;n del Estornino Pinto (\u003cem\u003eSturnus vulgaris\u003c/em\u003e) en un nido de Le\u0026ntilde;atero (\u003cem\u003eAnumbius annumbi\u003c/em\u003e) en el noreste de la provincia de Buenos Aires, Argentina. Nuestras Aves 59:13-14\u003c/li\u003e\n\u003cli\u003eFrei B, Nocera JJ, Fyles JW (2015) Interspecific competition and nest survival of the threatened Red-headed Woodpecker. J of Ornithol 156:743-753. https://doi.org/10.1007/s10336-015-1177-6 \u003c/li\u003e\n\u003cli\u003eFeare CJ (1984) The Starling. Oxford University Press, New York\u003c/li\u003e\n\u003cli\u003eGirini JM, Palacio FX, Del Huerto MDC, Kuzmanich N (2014) Selecci\u0026oacute;n de dormideros por el Estornino Pinto (\u003cem\u003eSturnus vulgaris\u003c/em\u003e) en La Plata, Buenos Aires, Argentina. El Hornero 29:23\u0026ndash;28\u003c/li\u003e\n\u003cli\u003eGomez De Silva H, Oliveras De Ita A, Legorreta RAM (2005) \u003cem\u003eSturnus vulgaris vulgaris\u003c/em\u003e. Vertebrados superiores ex\u0026oacute;ticos de M\u0026eacute;xico: diversidad, distribuci\u0026oacute;n y efectos potenciales. Instituto de Ecolog\u0026iacute;a, Universidad Nacional Aut\u0026oacute;noma de M\u0026eacute;xico, Ciudad de M\u0026eacute;xico\u003c/li\u003e\n\u003cli\u003eHowell SNG, Webb S (1995) A guide to the birds of Mexico and northern Central America. Oxford University Press, Oxford\u003c/li\u003e\n\u003cli\u003eIba\u0026ntilde;ez LM (2015) Invasi\u0026oacute;n del Estornino Pinto \u003cem\u003eSturnus vulgaris\u003c/em\u003e en el Noreste de la provincia de Buenos Aires: an\u0026aacute;lisis de la competencia con aves nativas y potencialidad como transmisor de par\u0026aacute;sitos. Doctoral thesis, Universidad Nacional de La Plata https://doi.org/10.35537/10915/45048 \u003c/li\u003e\n\u003cli\u003eIBGE (2019) Mapa de biomas e sistema costeiro marinho do Brasil - 1:250 000. Biblioteca IBGE. https://biblioteca.ibge.gov.br/index.php/biblioteca-catalogo?view=detalhes\u0026amp;id=2101676. Accessed 12 March 2025\u003c/li\u003e\n\u003cli\u003eIfran NR, Fiorini VD (2010) European Starling (\u003cem\u003eSturnus vulgaris\u003c/em\u003e): population density and interactions with native species in Buenos Aires urban parks. Ornitolog\u0026iacute;a Neotropical 21:507\u0026ndash;518\u003c/li\u003e\n\u003cli\u003eIsacch JP, Isacch J (2004) Estornino pinto (\u003cem\u003eSturnus vulgaris\u003c/em\u003e) en la ciudad de Mar del Plata (Buenos Aires, Argentina). Nuestras Aves 47:33. https://doi.org/10.56178/na.vi47.576 \u003c/li\u003e\n\u003cli\u003eKahle D, Wickham H (2013) ggmap: Spatial Visualization with ggplot2. The R J 5(1):144-161\u003c/li\u003e\n\u003cli\u003eKassambara A (2021) ggimage: Use Images in \u0026lsquo;ggplot2\u0026rsquo;. R package version 0.1.1\u003c/li\u003e\n\u003cli\u003eKlavins J, \u0026Aacute;lvarez D (2012) El Estornino Pinto (\u003cem\u003eSturnus vulgaris\u003c/em\u003e) en la provincia de C\u0026oacute;rdoba, Argentina. Nuestras Aves 57:27\u0026ndash;29\u003c/li\u003e\n\u003cli\u003eKnittle CE, Guarino JL (1976) Reducing a local population of starlings with nest-box traps. Bird Control Seminar Proceedings 7:65\u0026ndash;66\u003c/li\u003e\n\u003cli\u003eKoenig WD (2003) European starlings and their effect on native cavity‐nesting birds. Conserv Biol 17:1134\u0026ndash;1140. https://doi.org/10.1046/j.1523-1739.2003.02262.x \u003c/li\u003e\n\u003cli\u003eLinz GM, Homan HJ, Gaulker SM, Penry LB, Bleier WJ (2007) European starlings: a review of an invasive species with far-reaching impacts. Manag Vertebr Invasive Species 24:378-386\u003c/li\u003e\n\u003cli\u003eLockwood JL, Passy Y, Acubur R (2005) The role of propagule pressure in explaining species invasions. Trends in Ecol \u0026amp; Evol 20:223\u0026ndash;228\u003c/li\u003e\n\u003cli\u003eLowe S, Browne M, Boudjelas S, De Poorter M (2000) 100 of the world\u0026rsquo;s worst invasive alien species: a selection from the global invasive species database. Species Survival Commission, World Conservation Union (IUCN), Auckland\u003c/li\u003e\n\u003cli\u003eMazzulla J (2013) Primeros registros de Estornino Pinto \u003cem\u003eSturnus vulgaris\u003c/em\u003e (Linnaeus 1758) en Uruguay. Novid Ornitol\u0026oacute;gicas 3:1-17\u003c/li\u003e\n\u003cli\u003ePalacio FX, Iba\u0026ntilde;ez LM, Maragliano RE, Montalti D (2022) Uso del paisaje urbano por el Estornino Pinto (\u003cem\u003eSturnus vulgaris\u003c/em\u003e) durante las etapas reproductiva y no reproductiva. El Hornero 37(2):16\u003c/li\u003e\n\u003cli\u003ePebesma EJ, Bivand RS (2005) Classes and methods for spatial data in R. R News 5(2):2-11.\u003c/li\u003e\n\u003cli\u003eP\u0026eacute;rez J (1988) Estornino Pinto en la Capital Federal. Nuestras Aves 17:14.\u003c/li\u003e\n\u003cli\u003ePeris S, Soave G, Camperi A, Darrieu C, Aramburu R (2005) Range expansion of the European Starling \u003cem\u003eSturnus vulgaris\u003c/em\u003e in Argentina. Ardeola 52:359\u0026ndash;364.\u003c/li\u003e\n\u003cli\u003ePimentel D, Lach L, Zuniga R, Morrison D (2000) Environmental and economic costs of non indigenous species in the United States. BioScience 50:53-65. https://doi.org/10.1641/0006-3568(2000)050[0053:EAECON]2.3.CO;2 \u003c/li\u003e\n\u003cli\u003eRio Grande Do Sul (2024) Abertura da colheita da uva representa a cultura e a tradi\u0026ccedil;\u0026atilde;o ga\u0026uacute;cha. Secretaria da Agricultura, Pecu\u0026aacute;ria, Produ\u0026ccedil;\u0026atilde;o Sustent\u0026aacute;vel e Irriga\u0026ccedil;\u0026atilde;o. https://www.agricultura.rs.gov.br/aberta-colheita-da-uva-representa-a-cultura-e-tradicao-gaucha. Accessed 14 Jan 2025\u003c/li\u003e\n\u003cli\u003eRizzo F (2010) Utilizaci\u0026oacute;n de nidos de Hornero (\u003cem\u003eFurnarius rufus\u003c/em\u003e) por el Estornino Pinto \u003cem\u003e(Sturnus vulgaris\u003c/em\u003e). Nuestras Aves 55:33\u0026ndash;35\u003c/li\u003e\n\u003cli\u003eRuda Veja M (2004) Fotograf\u0026iacute;as curiosas y novedades de silvestres. Nuestras Aves 47:1.\u003c/li\u003e\n\u003cli\u003eR Core Team (2024). R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/\u003c/li\u003e\n\u003cli\u003eSchmidtutz C, Aguil\u0026aacute;n C (1988) Nidificaci\u0026oacute;n del estornino pinto. Nuestras Aves (17)\u003c/li\u003e\n\u003cli\u003eSilva FC, Da Pinto JM, M\u0026auml;der A, De Souza VAT (2017) First records of european starling \u003cem\u003eSturnus vulgaris\u003c/em\u003e in Brazil. Revista Bras de Ornitologia, 25(4):297-298. https://doi.org/10.1007/BF03544409 \u003c/li\u003e\n\u003cli\u003eSouza VA (2014) [WA1498234, \u003cem\u003eSturnus vulgaris\u003c/em\u003e Linnaeus, 1758]. Wiki Aves - A Enciclop\u0026eacute;dia das Aves do Brasil. http://www.wikiaves.com/1498234. Accessed 27 Jan 2025\u003c/li\u003e\n\u003cli\u003eWeber WJ (1979) Health hazards from pigeons, starlings and English sparrows: Diseases and parasites associated with pigeons, domestic animals, includes suggestions for bird control. Thomson Publications, New York\u003c/li\u003e\n\u003cli\u003eWickham H (2016) ggplot2- Elegant Graphics for Data Analysis 2nd Edition. Springer, New York. https://doi.org/10.18637/jss.v077.b02 \u003c/li\u003e\n\u003cli\u003eWing L (1943) Spread of the starling and European sparrow. The Auk 60:74-87\u003c/li\u003e\n\u003cli\u003eZanotti M (2013) Presencia del Estornino Pinto (\u003cem\u003eSturnus vulgaris\u003c/em\u003e) en la provincia de Mendoza, Argentina. Nuestras Aves 58:5\u0026ndash;7\u003c/li\u003e\n\u003cli\u003eZufiaurre E, Abba A, Bilenca D, Codesido M (2016) Role of landscape elements on recent distributional expansion of European starlings (\u003cem\u003eSturnus vulgaris\u003c/em\u003e) in agroecosystems of the Pampas, Argentina. Wilson J of Ornithol 128:306\u0026ndash;313. https://doi.org/10.1676/wils-128-02-306-313.1\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"
[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Sturnidae, exotic bird, invasive species, Pampa biome","lastPublishedDoi":"10.21203/rs.3.rs-7574299/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7574299/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eWe conducted a survey of the records of \u003cem\u003eSturnus vulgaris\u003c/em\u003e (the European starling) in Brazil over the 10 years following its first documented occurrence in the country. The records were gathered from citizen science platforms, field data, and reports from researchers and birdwatchers. A total of 2,618 individuals were recorded across 219 occurrences, 214 of which took place in Rio Grande do Sul, with five others distributed among the states of S\u0026atilde;o Paulo, Rio de Janeiro, and Minas Gerais. In just one decade, the species has already occupied a considerable portion of the Brazilian Pampa, especially in the southernmost regions near the border with Uruguay, with 97.7% of the records concentrated in Rio Grande do Sul. We identified 55 nesting sites, most of which were located in cavities previously excavated by woodpeckers (Picidae), primarily in trees of the genus \u003cem\u003eEucalyptus\u003c/em\u003e. Breeding begins in September, peaks between October and November, and extends until the end of December. The data from this study indicate a high degree of behavioral plasticity in the species, facilitating its rapid adaptation and expansion within the Pampa biome.\u003c/p\u003e","manuscriptTitle":"Ten years of the European Starling (Sturnus vulgaris) in Brazil: range expansion and reproductive insights","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-11-06 07:16:10","doi":"10.21203/rs.3.rs-7574299/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"
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