REFERENCES
Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ (1990) Basic Local Alignment Search Tool. Journal of Molecular Biology ,215 , 403–410.
Ashenden T, Rooke AC, Fox MG (2017) Boldness and dispersal tendency of native and invasive pumpkinseed (Lepomis gibbosus ): is spatial sorting creating superior invaders? Aquatic Invasions , 12 , 311-320.
Avalos A, Pan H, Li C et al. (2017) A soft selective sweep during rapid evolution of gentle behaviour in an Africanized honeybee.Nature Communications , 1–9.
Baltazar-Soares M, Blanchet S, Cote J, Tarkan AS, Záhorská, Gozlan RE, Eizaguirre C (2019) Genomic footprints of a biological invasion: introduction from Asia and dispersal in Europe of the topmouth gudgeon (Pseudorasbora parva ). Molecular Ecology , 29 , 71-85.
Bay RA, Harrigan RJ, Le Underwood V et al. (2018) Genomic signals of selection predict climate-driven population declines in a migratory bird. Science , 359 , 83–86.
Berthouly-Salazar C, van Rensburg BJ, Le Roux JJ, van Vuuren BJ, Hui C (2012) Spatial sorting drives morphological variation in the invasive bird, Acridotheris tristis . PLoS ONE , 7 , e38145.
Bitton P-P, Graham BA (2014) Change in wing morphology of the European starling during and after colonization of North America. Journal of Zoology , 295 , 254–260.
Briski E, Chan FT, Darling JA, Lauringson V, MacIsaac HJ, Zhan A, Bailey SA (2018) Beyond propagule pressure: importance of selection during the transport stage of biological invasions. Frontiers in Ecology and the Environment , 16 , 345-353.
Brown GP, Phillips BL, Shine R (2014) The straight and narrow path: the evolution of straight-line dispersal at a cane toad invasion front.Proceedings of the Royal Society B , 281 , 20141385.
Burtt HE, Giltz ML (1977) Seasonal directional patterns of movements and migrations of Starlings and Blackbirds in North America.Bird-Banding , 48 , 259-271.
Cabe PR (1998) The effects of founding bottlenecks on genetic variation in the European starling (Sturnus vulgaris ) in North America.Heredity , 80 , 519–525.
Cabe PR (1999) Dispersal and population structure in the European Starling. The Condor , 101 , 451–454.
Capblancq T, Luu K, Blum MGB, Bazin E (2018) Evaluation of redundancy analysis to identify signatures of local adaptation. Molecular Ecology Resources , 185 , 587–11.
Cardador L, Blackburn TM (2019) Human-habitat associations in the native distrbutions of alien bird species. J Applied Ecology ,56 , 1189-1199.
Cardilini APA, Stuart KC, Cassey P, Richardson MF, Sherwin W, Rollins LA, Sherman CDH (2020) Signatures of selection in a recent invasion reveals adaptive divergence in a highly vagile invasive species. bioRxiv, doi: https://doi.org/10.1101/643569.
Cariou M, Duret L, Charlat S (2016) How and how much does RAD-seq bias genetic diversity estimates? BMC Evolutionary Biology , 1–8.
Catchen J, Hohenlohe PA, Bassham S, Amores A, Cresko WA (2013) Stacks: an analysis tool set for population genomics. Molecular Ecology ,22 , 3124–3140.
Cohen, TM, McKinney, M, Kark S, Dor R (2019) Global invasion in progress: modeling the past, current, and potential global distribution of the common myna. Biological Invasions , 21 , 1295-1309.
Colautti RI, Lau JA (2015) Contemporary evolution during invasion: evidence for differentiation, natural selection, and local adaptation.Molecular Ecology , 24 , 1999–2017.
Danecek P, Auton A, Abecasis G et al. (2011) The variant call format and VCFtools. Bioinformatics , 27 , 2156–2158.
de Villemereuil P, Gaggiotti OE (2015) A new FST‐based method to uncover local adaptation using environmental variables. Methods in Ecology and Evolution , 6 , 1248–1258.
Dlugosch KM, Parker IM (2008) Founding events in species invasions: genetic variation, adaptive evolution, and the role of multiple introductions. Molecular Ecology , 17 , 431–449.
Dlugosch KM, Anderson SR, Braasch J, Cang FA, Gillette HD (2015) The devil is in the details: genetic variation in introduced populations and its contributions to invasion. Molecular Ecology , 24 , 2095–2111.
Dolbeer RA (1982) Migration patterns for age and sex classes of Blackbirds and Starlings. Journal of Field Ornithology ,53 , 28-46.
Duckworth RA (2008) Adaptive Dispersal Strategies and the Dynamics of a Range Expansion. The American Naturalist , 172 , S4–S17.
Earl DA, vonHoldt BM (2011) STRUCTURE HARVESTER: a website and program for visualizing STRUCTURE output and implementing the Evanno method.Conservation Genetics Resources , 4 , 359–361.
Edelaar P, Bolnick DI (2012) Non-random gene flow: an underappreciated force in evolution and ecology. Trends in Ecology & Evolution ,27 , 659–665.
Edelaar P, Roques S, Hobson EA, Da Silva AG, Avery ML, Russello MA, Senar JC, Wright TF, Carrete M, Tella JL (2015) Shared genetic diversity across the global invasive range of the monk parakeet suggests a common restricted geographic origin and the possibility of convergent selection. Molecular Ecology , 24 , 2164-2716.
Estoup A, Ravigné V, Hufbauer R et al. (2016) Is There a Genetic Paradox of Biological Invasion? Annual Review of Ecology, Evolution, and Systematics , 47 , 51–72.
Excoffier L, Dupanloup I, Huerta-Sánchez E, Sousa VC, Foll M (2013) Robust demographic inference from genomic and SNP data. PLOS Genetics, 9 , e1003905.
Facon B, Hufbauer RA, Tayeh A et al. (2011) Inbreeding Depression Is Purged in the Invasive Insect Harmonia axyridis. Current Biology , 21 , 424–427.
Fick SE, Hijmans RJ (2017) WorldClim 2: New 1-km spatial resolution climate surfaces for global land areas . International Journal of Climatology .
Forbush EH (1915) The starling. Boston, Wright & Potter Printing Co.
Forester BR, Lasky JR, Wagner HH, Urban DL (2018) Comparing methods for detecting multilocus adaptation with multivariate genotype-environment associations. Molecular Ecology , 27 , 2215–2233.
Frichot E, François O (2015) LEA: An R package for landscape and ecological association studies (B O’Meara, Ed,). Methods in Ecology and Evolution , 6 , 925–929.
Frichot E, Schoville SD, Bouchard G, François O (2013) Testing for Associations between Loci and Environmental Gradients Using Latent Factor Mixed Models. Molecular Biology and Evolution ,30 , 1687–1699.
Garant D, Forde SE, Hendry AP (2007) The multifarious effects of dispersal and gene flow on contemporary adaptation. Functional Ecology , 21 , 434–443.
Gralka M, Hallatschek O (2019) Environmental heterogeneity can tip the population genetics of range expansions. eLife , 8 , e44359.
Guillot G, Rousset F (2013) Dismantling the Mantel tests (L Harmon, Ed,). Methods in Ecology and Evolution , 4 , 336–344.
Günther T, Coop GC (2013) Robust identification of local adaptation from allele frequencies. Genetics Soc America , 195 , 205-220.
Hijmans RJ, van Etten J (2012) raster: Geographic analysis and modeling with raster data. R package version 2.0-12.
Hoban S, Kelley JL, Lotterhos KE et al. (2016) Finding the Genomic Basis of Local Adaptation: Pitfalls, Practical Solutions, and Future Directions. The American Naturalist , 188 , 379–397.
Hufbauer RA, Facon B, Ravigné V et al. (2011) Anthropogenically induced adaptation to invade (AIAI): contemporary adaptation to human-altered habitats within the native range can promote invasions.Evolutionary Applications , 5 , 89–101.
Hui C, Roura-Pascual N, Brotons L, Robinson RA, Evans KL (2012) Flexible dispersal strategies in native and non-native ranges: environmental quality and the “good-stay, bad-disperse” rule. Ecography ,35 , 1024–1032.
Jacob S, Legrand D, Chaine AS et al. (2017) Gene flow favours local adaptation under habitat choice in ciliate microcosms.Nature Ecology & Evolution , 1–4.
Jakobsson M, Rosenberg NA (2007) CLUMPP: a cluster matching and permutation program for dealing with label switching and multimodality in analysis of population structure. Bioinformatics , 23 , 1801–1806.
Jernelov A (2017) Starlings in North America. In: The Long-Term Fate of Invasive Species , pp. 137–149. Jarpas.
Jombart T (2008) adegenet: a R package for the multivariate analysis of genetic markers. Bioinformatics , 24 , 1403–1405.
Kalmbach, ER, and IN Gabrielson (1921) Economic Value of the Starling in the United States USDA Bulletin , 868 , 1-59.
Kamvar ZN, Tabima JF, Grünwald NJ (2014) Poppr: an R package for genetic analysis of populations with clonal, partially clonal, and/or sexual reproduction. PeerJ , 2 , e281–14.
Kessel B (1953) Distribution and migration of the European Starling in North America. The Condor , 55 , 49–67.
Koenig W (2003) European Starlings and Their Effect on Native Cavity-Nesting Birds. Conservation Biology , 17 , 1134–1140.
Langmead B, Salzberg SL (2012) Fast gapped-read alignment with Bowtie 2.Nature Methods , 9 , 357–359.
Lenormand T (2002) Gene flow and the limits to natural selection.17 , 183–189.
Lescak EA, Bassham SL, Catchen J et al. (2015) Evolution of stickleback in 50 years on earthquake-uplifted islands.Proceedings of the National Academy of Sciences , 112 , E7204–E7212.
Leydet KP, Grupstra CGB, Coma R, Ribes M, Hellberg ME (2018) Host-targeted RAD-Seq reveals genetic changes in the coral Oculina patagonica associated with range expansion along the Spanish Mediterranean coast. Mol Ecol , 27 , 2529-2543.
Linck EB, Battey CJ (2017) Minor allele frequency thresholds strongly affect population structure inference with genomic datasets.bioRxiv .
Linz GM, Homan HJ, Gaulker SM, Penry LB, Bleier WJ (2007) European starlings: a review of an invasive species with far-reaching impacts .
Liebl AL, Schrey AW, Richards, CL, Martin LB (2013) Patterns of DNA methylation throughout a range expansion of an introduced songbird.Integ Comp Bio , 53 , 351-358.
Liu X, Fu Y-X (2015) Exploring population size changes using SNP frequency spectra. Nature Genetics , 47 , 555–559.
Lombaert E, Estoup A, Facon B et al. (2014) Rapid increase in dispersal during range expansion in the invasive ladybird Harmonia axyridis . Journal of Evolutionary Biology , 27 , 508–517.
Marin P, Genitoni J, Barloy D, Maury S, Gibert P, Ghalambor CK, Vieira C (2019) Biological invasion: The influence of the hidden side of the (epi)genome. Functional Ecology, 34, 385-400.
Malinsky M, Trucchi E, Lawson D, Falush D (2016) RADpainter and fineRADstructure: population inference from RADseq data. 1–6.
Meirmans PG (2015) Seven common mistakes in population genetics and how to avoid them. Molecular Ecology , 24 , 3223-3231.
Messer PW, Petrov DA (2013) Population genomics of rapid adaptation by soft selective sweeps. Trends in Ecology & Evolution ,28 , 659–669.
Mi H, Dong Q, Muruganujan A et al. (2009) PANTHER version 7: improved phylogenetic trees, orthologs and collaboration with the Gene Ontology Consortium. Nucleic Acids Research , 38 , D204–D210.
Ochocki BM, Miller TEX (2017) Rapid evolution of dispersal ability makes biological invasions faster and more variable. Nature Communications , 8 , 1–8.
Oksanen J, Blanchet FG, Friendly M, Kindt R, Legendre P, McGlinn D, Minchin PR, O’Hara RB, Simpson GL, Solymos P, Stevens MHH, Szoecs E, Wagner H (2019). vegan: Community Ecology Package. R package version 2.5-6. https://CRAN.R-project.org/package=vegan
Peterson BK, Weber JN, Kay EH, Fisher HS, Hoekstra HE (2012) Double Digest RADseq: An Inexpensive Method for De Novo SNP Discovery and Genotyping in Model and Non-Model Species (L Orlando, Ed,). PLoS ONE , 7 , e37135–11.
Petkova D, Novembre J, Stephens M (2015) Visualizing spatial population structure with estimated effective migration surfaces. Nature Reviews Neuroscience , 48 , 94–100.
Pfeifer SP, Laurent S, Sousa VC et al. (2018) The Evolutionary History of Nebraska Deer Mice: Local Adaptation in the Face of Strong Gene Flow (N Singh, Ed,). Molecular Biology and Evolution ,35 , 792–806.
Phair DJ, Le Roux J, Berthouly-Salazar C et al (2018) Context-dependent spatial sorting of dispersal-related traits in the invasive starlings (Sturnus vulgaris) of South Africa and Australia. bioRxiv,342451.
Phillips BL, Perkins TA (2019) Spatial sorting as the spatial analogue of natural selection. Theoretical Ecology , 12 , 155-163.
Prentis PJ, Wilson JRU, Dormontt EE, Richardson DM, Lowe AJ (2008) Adaptive evolution in invasive species. Trends in Plant Science ,13 , 288–294.
Pritchard JK, Stephens M, Donnelly P (2000) Inference of Population Structure Using Multilocus Genotype Data. Genetics , 155 , 945–959.
Quinlan AR, Hall IM (2010) BEDTools: a flexible suite of utilities for comparing genomic features. Bioinformatics , 26 , 841–842.
R Core Team (2019). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/.
Ravinet M, Elgin TO, Trier C, Aliabadian M, Gavrilov A, Sætre, G-P (2018) Signatures of human-commensalism in the house sparrow genome.Proceedings of the Royal Society B , 285 , 20181246.
Rius M, Darling JA (2014) How important is intraspecific genetic admixture to the success of colonising populations? Trends in Ecology & Evolution , 29 , 233–242.
Rollins LA, Richardson MF, Shine R (2015) A genetic perspective on rapid evolution in cane toads ( Rhinella marina). Molecular Ecology ,24 , 2264–2276.
Rollins LA, Woolnough AP, Fanson BG et al. (2016) Selection on Mitochondrial Variants Occurs between and within Individuals in an Expanding Invasion. Molecular Biology and Evolution , 33 , 995–1007.
Rollins LA, Woolnough AP, Sinclair R, Mooney NJ, Sherwin WB (2011) Mitochondrial DNA offers unique insights into invasion history of the common starling. Molecular Ecology , 20 , 2307–2317.
Rollins LA, Woolnough AP, Wilton AN, Sinclair R, Sherwin WB (2009) Invasive species can’t cover their tracks: using microsatellites to assist management of starling (Sturnus vulgaris ) populations in Western Australia. Molecular Ecology , 18 , 1560–1573.
Rosenberg NA (2003) distruct: a program for the graphical display of population structure. Molecular Ecology Notes , 4 , 137–138.
Schlaepfer DR, Glättli M, Fischer M, van Kleunen M (2009) A multi-species experiment in their native range indicates pre-adaptation of invasive alien plant species. New Phytologist , 185 , 1087–1099.
Schmid-Hempel P, Schmid-Hempel R, Brunner PC, Seeman OD, Allen GR (2007) Invasion success of the bumblebee, Bombus terrestris, despite a drastic genetic bottleneck. Heredity , 99 , 414–422.
Schrieber K, Lachmuth L (2017) The Genetic Paradox of Invasions revisited: the potential role of inbreeding X environment interactions in invasion success. Biol Rev , 92 , 939-952.
Schweizer RM, Robinson J, Harrigan R et al. (2015) Targeted capture and resequencing of 1040 genes reveal environmentally driven functional variation in grey wolves. Molecular Ecology ,25 , 357–379.
Sexton JP, Hangartner SB, Hoffmann AA (2014) Genetic isolation by environment or distance: which pattern of gene flow is most common?Evolution , 68 , 1–15.
Sheldon EL, Schrey A, Andrew SC, Ragsdale A, Griffith SC (2018) Epigenetic and genetic variation among three separate introductions of the house sparrow (Passer domesticus ) into Australia. R. Soc. Open Sci , 5 , 172185.
Szűcs M, Vahsen ML, Melbourne BA et al. (2017) Rapid adaptive evolution in novel environments acts as an architect of population range expansion. Proceedings of the National Academy of Sciences ,114 , 13501–13506.
Thomas PD, Campbell MJ, Kejariwal A et al. (2003) PANTHER: A Library of Protein Families and Subfamilies Indexed by Function.Genome Research , 13 , 2129–2141.
Thrasher DJ, Butcher BG, Campagna L, Webster MS, Lovette IJ (2017) Double-digest RAD sequencing outperforms microsatellite loci at assigning paternity and estimating relatedness: a proof of concept in a highly promiscuous bird. 1–34.
Tigano A, Friesen VL (2016) Genomics of local adaptation with gene flow.Molecular Ecology , 25 , 2144–2164.
Tsutsui ND, Suarez AV, Holway DA, Case TJ (2000) Reduced genetic variation and the success of an invasive species. Proceedings of the National Academy of Sciences , 97 , 5948–5953.
Uller T, Leimu R (2011) Founder events predict changes in genetic diversity during human-mediated range expansions. Global Change Biology , 17 , 3478–3485.
Werner SJ, Fischer JW, Hobson KA (in press) Multi-isotopic (δ2H, δ13C, δ15N) tracing of molt origin for European starlings associated with U.S. dairies and feedlots. PLoS One .
Willoughby JR, Harder AM, Tennessen JA, Scribner KT, Christie MR (2018) Rapid genetic adaptation to a novel environment despite a genome‐wide reduction in genetic diversity. Molecular Ecology , 17 , 675.
Yeaman S, Whitlock MC (2011) The genetic architecture of adaptation under migration-selection balance. Evolution , 65 , 1897–1911.
Zheng X, Levine D, Shen J et al. (2012) A high-performance computing toolset for relatedness and principal component analysis of SNP data. Bioinformatics , 28 , 3326–3328.
Figure Captions
Table 1. Population genetic summary statistics. For each sampling location, the table shows the number of private alleles, observed and expected heterozygosity, nucleotide diversity, and the inbreeding coefficient FIS.
Table 2. Differentiation and movement among sampling locations. This table shows the percentage of birds assigned to each sampling location from every origin. Note that these values are directional: although 12 starlings that were collected in AZ originated in KS, only 2 starlings collected in KS originated in AZ. The black diagonal indicates the percentage of birds assigned to that collection state according to a discriminant function analysis of molt-origin presented in Werner et al. For example, 28% of birds collected in Arizona originated in that location. FST among locations is presented above the diagonal, and darker gray colors indicate higher FST.
Figure 1. Population structure. A) Principal components analysis on 6,287 SNPs, explaining 1.03-1.08% of genetic variation observed. B) STRUCTURE analyses with K=2 and K=3 (best supported). C) Significance testing of hierarchical AMOVAs: the histogram shows expected variance components based on 999 simulations, and the black diamond is the observed variance component. D) AMOVA results.
Figure 2. Evidence for incipient local adaptation. Redundancy analyses indicate that 191 SNPs (small gray points) are associated with bioclimatic predictors (vectors). BIO1: mean annual temperature; BIO7: temperature annual range; BIO12: annual precipitation; BIO16: precipitation of wettest quarter.