The speed of change in climate and land cover is associated with the speed of biodiversity changes in avian assemblages of the United States

sition of the bird assemblages and the rate of change of the environment in different regions of the world. In any case, our results highlight the importance of considering multiple environmental factors and ecoregion-specific analysis in com- munity ecology since threats to species assemblages can arise from changes in climate or LULC independently, or their combined effect [15], and vary per region [18,19]. Conclusions Changes in climate and/or LULC are driving the processes of community assembly and breakdown more than just driving the pattern of species richness and these assemblies and breakdowns can occur near one another within the same ecoregion. The more obvious changes are occurring in high altitude and high latitude ecoregions. For some ecoregions, components of total assemblage dissimilarity are responding to the environment in opposite ways. For all ecoregions, LULC was a stronger predictor of change in biodiversity metrics than was climate. Assemblage dissimi- larity was better predicted (by the environment) than was species richness, despite the fact that species richness is the better studied phenomenon. It is only through examining trends in multiple biodiversity indices, meant to capture multiple levels of action (species richness versus assemblage dissimilarity) that we would be able to observe the complex and changing interrelationships between biodiversity (species richness and assemblage dissimilarity) and the environment (climate and LULC).

PLOS One | https://doi.org/10.1371/journal.pone.0330153 August 29, 2025 16 / 20 Supporting information S1 File. Combined file containing the supporting information associated with this article. (DOCX) Acknowledgments We would like to acknowledge all the volunteers who have contributed to citizen science projects such as the North American Breeding Bird Survey. We would like to thank Dr. Shannon R. Curley and Dr. Richard R. Veit for their insightful comments and suggestions that improved the paper. Author contributions Conceptualization: Marlen Acosta Alamo, Lisa L. Manne. Data curation: Marlen Acosta Alamo. Formal analysis: Marlen Acosta Alamo. Funding acquisition: Lisa L. Manne. Investigation: Marlen Acosta Alamo. Methodology: Marlen Acosta Alamo, Lisa L. Manne. Project administration: Lisa L. Manne. Supervision: Lisa L. Manne. Visualization: Marlen Acosta Alamo. Writing – original draft: Marlen Acosta Alamo. Writing – review & editing: Marlen Acosta Alamo, Lisa L. Manne. References 1. Davey CM, Chamberlain DE, Newson SE, Noble DG, Johnston A. Rise of the generalists: evidence for climate driven homogenization in avian communities. Global Ecol Biogeo. 2011;21(5):568–78. https://doi.org/10.1111/j.1466-8238.2011.00693.x 2. Magurran AE, Dornelas M, Moyes F, Gotelli NJ, McGill B. Rapid biotic homogenization of marine fish assemblages. Nat Commun. 2015;6:8405. https://doi.org/10.1038/ncomms9405 PMID: 26400102 3. McKinney M, Lockwood J. Biotic homogenization: a few winners replacing many losers in the next mass extinction. Trends Ecol Evol.

1999;14(11):450–3. https://doi.org/10.1016/s0169-5347(99)01679-1 PMID: 10511724 4. Savage J, Vellend M. Elevational shifts, biotic homogenization and time lags in vegetation change during 40 years of climate warming. Ecography. 2014;38(6):546–55. https://doi.org/10.1111/ecog.01131 5. White HJ, Montgomery WI, Storchová L, Hořák D, Lennon JJ. Does functional homogenization accompany taxonomic homogenization of British birds and how do biotic factors and climate affect these processes?. Ecol Evol. 2018;8(15):7365–77. https://doi.org/10.1002/ece3.4267 PMID: 30151156 6. Emslie SD. Avian community, climate, and sea-level changes in the plio-pleistocene of the Florida Peninsula. Ornithological Monographs. 1998;(50):1–113. https://doi.org/10.2307/40166707 7. Webb T. Pollen records of late Quaternary vegetation change: plant community rearrangements and evolutionary implications. 1995. 221–32. 8. Loarie SR, Duffy PB, Hamilton H, Asner GP, Field CB, Ackerly DD. The velocity of climate change. Nature. 2009;462(7276):1052–5. https://doi. org/10.1038/nature08649 PMID: 20033047 9. García‐Valdés R, Zavala MA, Araújo MB, Purves DW. Chasing a moving target: projecting climate change‐induced shifts in non‐equilibrial tree species distributions. J Ecol. 2013;101(2):441–53. https://doi.org/10.1111/1365-2745.12049 10. Bellard C, Leclerc C, Courchamp F. Combined impacts of global changes on biodiversity across the USA. Sci Rep. 2015;5:11828. https://doi. org/10.1038/srep11828 PMID: 26149694

PLOS One | https://doi.org/10.1371/journal.pone.0330153 August 29, 2025 17 / 20 11. Stegner MA, Spanbauer TL. North American pollen records provide evidence for macroscale ecological changes in the Anthropocene. Proc Natl Acad Sci U S A. 2023;120(43):e2306815120. https://doi.org/10.1073/pnas.2306815120 PMID: 37844232 12. Urban MC. Climate change. Accelerating extinction risk from climate change. Science. 2015;348(6234):571–3. https://doi.org/10.1126/science. aaa4984 PMID: 25931559 13. Newbold T, Hudson LN, Hill SLL, Contu S, Lysenko I, Senior RA, et al. Global effects of land use on local terrestrial biodiversity. Nature. 2015;520(7545):45–50. https://doi.org/10.1038/nature14324 PMID: 25832402 14. Newbold T, Hudson LN, Arnell AP, Contu S, De Palma A, Ferrier S, et al. Has land use pushed terrestrial biodiversity beyond the planetary bound- ary? A global assessment. Science. 2016;353(6296):288–91. https://doi.org/10.1126/science.aaf2201 PMID: 27418509 15. Caro T, Rowe Z, Berger J, Wholey P, Dobson A. An inconvenient misconception: climate change is not the principal driver of biodiversity loss. Con- serv Lett. 2022;15(3). https://doi.org/10.1111/conl.12868 16. Jaureguiberry P, Titeux N, Wiemers M, Bowler DE, Coscieme L, Golden AS, et al. The direct drivers of recent global anthropogenic biodiversity loss. Sci Adv. 2022;8(45):eabm9982. https://doi.org/10.1126/sciadv.abm9982 PMID: 36351024 17. Newbold T. Future effects of climate and land-use change on terrestrial vertebrate community diversity under different scenarios. Proc Biol Sci.

24. Ferger SW, Peters MK, Appelhans T, Detsch F, Hemp A, Nauss T, et al. Synergistic effects of climate and land use on avian beta‐diversity. Diversity Distributions. 2017;23(11):1246–55. https://doi.org/10.1111/ddi.12615 25. Ganuza C, Redlich S, Uhler J, Tobisch C, Rojas-Botero S, Peters MK, et al. Interactive effects of climate and land use on pollinator diversity differ among taxa and scales. Sci Adv. 2022;8(18):eabm9359. https://doi.org/10.1126/sciadv.abm9359 PMID: 35544641 26. Gaüzère P, Barbaro L, Calatayud F, Princé K, Devictor V, Raison L, et al. Long-term effects of combined land-use and climate changes on local bird communities in mosaic agricultural landscapes. Agricul Ecosyst Environ. 2020;289:106722. https://doi.org/10.1016/j.agee.2019.106722 27. Karp DS, Frishkoff LO, Echeverri A, Zook J, Juárez P, Chan KMA. Agriculture erases climate-driven β-diversity in Neotropical bird communities. Glob Chang Biol. 2018;24(1):338–49. https://doi.org/10.1111/gcb.13821 PMID: 28833924 28. MacLean SA, Rios Dominguez AF, de Valpine P, Beissinger SR. A century of climate and land-use change cause species turnover without loss of beta diversity in California’s central valley. Glob Chang Biol. 2018;24(12):5882–94. https://doi.org/10.1111/gcb.14458 PMID: 30267548 29. Oliver I, Dorrough J, Doherty H, Andrew NR. Additive and synergistic effects of land cover, land use and climate on insect biodiversity. Landscape Ecol. 2016;31(10):2415–31. https://doi.org/10.1007/s10980-016-0411-9

30. Oliver TH, Gillings S, Pearce-Higgins JW, Brereton T, Crick HQP, Duffield SJ, et al. Large extents of intensive land use limit community reorganiza- tion during climate warming. Glob Chang Biol. 2017;23(6):2272–83. https://doi.org/10.1111/gcb.13587 PMID: 28073167 31. Wayman JP, Sadler JP, Pugh TAM, Martin TE, Tobias JA, Matthews TJ. Assessing taxonomic and functional change in British breeding bird assem- blages over time. Global Ecol Biogeogr. 2022;31(5):925–39. https://doi.org/10.1111/geb.13468 32. Zerbo I, Bernhardt-Römermann M, Ouédraogo O, Hahn K, Thiombiano A. Effects of climate and land use on herbaceous species richness and vegetation composition in West African Savanna ecosystems. J Botany. 2016;2016:1–11. https://doi.org/10.1155/2016/9523685 33. Hillebrand H, Matthiessen B. Biodiversity in a complex world: consolidation and progress in functional biodiversity research. Ecol Lett. 2009;12(12):1405–19. https://doi.org/10.1111/j.1461-0248.2009.01388.x PMID: 19849711 34. Karp DS, Rominger AJ, Zook J, Ranganathan J, Ehrlich PR, Daily GC. Intensive agriculture erodes β-diversity at large scales. Ecol Lett. 2012;15(9):963–70. https://doi.org/10.1111/j.1461-0248.2012.01815.x PMID: 22727063 35. Pecl GT, Araújo MB, Bell JD, Blanchard J, Bonebrake TC, Chen I-C, et al. Biodiversity redistribution under climate change: impacts on ecosystems and human well-being. Science. 2017;355(6332):eaai9214. https://doi.org/10.1126/science.aai9214 PMID: 28360268

36. Scheffers BR, De Meester L, Bridge TCL, Hoffmann AA, Pandolfi JM, Corlett RT, et al. The broad footprint of climate change from genes to biomes to people. Science. 2016;354(6313):aaf7671. https://doi.org/10.1126/science.aaf7671 PMID: 27846577 37. Devictor V, van Swaay C, Brereton T, Brotons L, Chamberlain D, Heliölä J, et al. Differences in the climatic debts of birds and butterflies at a conti- nental scale. Nature Clim Change. 2012;2(2):121–4. https://doi.org/10.1038/nclimate1347

PLOS One | https://doi.org/10.1371/journal.pone.0330153 August 29, 2025 18 / 20 38. Huang M, Piao S, Janssens IA, Zhu Z, Wang T, Wu D, et al. Velocity of change in vegetation productivity over northern high latitudes. Nat Ecol Evol. 2017;1(11):1649–54. https://doi.org/10.1038/s41559-017-0328-y PMID: 28970570 39. Pinsky ML, Worm B, Fogarty MJ, Sarmiento JL, Levin SA. Marine taxa track local climate velocities. Science. 2013;341(6151):1239–42. https://doi. org/10.1126/science.1239352 PMID: 24031017 40. Asamoah EF, Beaumont LJ, Maina JM. Climate and land-use changes reduce the benefits of terrestrial protected areas. Nat Clim Chang. 2021;11(12):1105–10. https://doi.org/10.1038/s41558-021-01223-2 41. Ordonez A, Martinuzzi S, Radeloff VC, Williams JW. Combined speeds of climate and land-use change of the conterminous US until 2050. Nature Clim Change. 2014;4(9):811–6. https://doi.org/10.1038/nclimate2337 42. Acosta Alamo M, Manne LL. Rapid rates of change in multiple biodiversity measures in breeding Avian assemblages. Diversity. 2025;17(2):127. https://doi.org/10.3390/d17020127 43. Baselga A, Orme CDL. betapart: an R package for the study of beta diversity. Methods Ecol Evol. 2012;3(5):808–12. https://doi. org/10.1111/j.2041-210x.2012.00224.x 44. Burgess MD, Smith KW, Evans KL, Leech D, Pearce-Higgins JW, Branston CJ, et al. Tritrophic phenological match-mismatch in space and time. Nat Ecol Evol. 2018;2(6):970–5. https://doi.org/10.1038/s41559-018-0543-1 PMID: 29686235

45. Kellermann JL, van Riper C 3rd. Detecting mismatches of bird migration stopover and tree phenology in response to changing climate. Oecologia. 2015;178(4):1227–38. https://doi.org/10.1007/s00442-015-3293-7 PMID: 25822114 46. Blois JL, Zarnetske PL, Fitzpatrick MC, Finnegan S. Climate change and the past, present, and future of biotic interactions. Science. 2013;341(6145):499–504. https://doi.org/10.1126/science.1237184 PMID: 23908227 47. Ziolkowski DJJr, Lutmerding M, Aponte VI, Hudson MR. North American breeding bird survey dataset 1966–2021: U.S. geological survey data release. Laurel, MD, USA: U.S. Geological Survey; 2022. https://doi.org/10.5066/P97WAZE5 48. Aiello‐Lammens ME, Boria RA, Radosavljevic A, Vilela B, Anderson RP. spThin: an R package for spatial thinning of species occurrence records for use in ecological niche models. Ecography. 2015;38(5):541–5. https://doi.org/10.1111/ecog.01132 49. Daly C, Taylor GH, Gibson WP, Parzybok TW, Johnson GL, Pasteris PA. High-quality spatial climate data sets for the United States and beyond. Trans ASAE. 2000;43(6):1957–62. https://doi.org/10.13031/2013.3101 50. Jeffree EP, Jeffree CE. Temperature and the biogeographical distributions of species. Funct Ecol. 1994;8(5):640. https://doi.org/10.2307/2389927 51. Root T. Energy constraints on avian distributions and abundances. Ecology. 1988;69(2):330–9. https://doi.org/10.2307/1940431 52. Sinnott EA, Papeş M, O’Connell TJ. Variable precipitation leads to dynamic range limits of forest songbirds at a forest-grassland ecotone. Ecol

Evol. 2021;11(16):11123–33. https://doi.org/10.1002/ece3.7899 PMID: 34429907 53. Sohl TL. Modeled historical land use and land cover for the conterminous United States: 1938-1992: U.S. geological survey data release. 2018. Accessed 02 January 2023. https://doi.org/10.5066/F7KK99RR 54. Sohl TL. Conterminous United States land cover projections - 1992 to 2100. 2018. https://doi.org/10.5066/P95AK9HP 55. United States Department of the Interior. North America elevation. U.S. geological survey. 2007. Accessed 2023 February 15. http://www.cec.org/ north-american-environmental-atlas/elevation-2007/ 56. Belmaker J, Jetz W. Cross-scale variation in species richness-environment associations. Global Ecol Biogeo. 2010;20(3):464–74. https://doi. org/10.1111/j.1466-8238.2010.00615.x 57. Wood SN. Fast stable restricted maximum likelihood and marginal likelihood estimation of semiparametric generalized linear models. J Royal Stat Soc Series B: Stat Methodol. 2010;73(1):3–36. https://doi.org/10.1111/j.1467-9868.2010.00749.x 58. Wood SN. Generalized additive models: An Introduction with R. Chapman and Hall/CRC; 2017. https://doi.org/10.1201/9781315370279 59. Whittaker RH. Vegetation of the Siskiyou mountains, Oregon and California. Ecol Monographs. 1960;30(3):279–338. https://doi. org/10.2307/1943563 60. Flores O, Seoane J, Hevia V, Azcárate FM. Spatial patterns of species richness and nestedness in ant assemblages along an elevational gradient in a Mediterranean mountain range. PLoS One. 2018;13(12):e0204787. https://doi.org/10.1371/journal.pone.0204787 PMID: 30566476

61. Gordó-Vilaseca C, Stephenson F, Coll M, Lavin C, Costello MJ. Three decades of increasing fish biodiversity across the northeast Atlantic and the Arctic Ocean. Proc Natl Acad Sci U S A. 2023;120(4):e2120869120. https://doi.org/10.1073/pnas.2120869120 PMID: 36656855 62. Curley SR, Manne LL, Ramírez‐Garofalo JR, Veit RR. Evaluating compositional changes in the avian communities of eastern North America using temperature and precipitation indices. J Biogeo. 2022;49(4):739–52. https://doi.org/10.1111/jbi.14340 63. Tayleur CM, Devictor V, Gaüzère P, Jonzén N, Smith HG, Lindström Å. Regional variation in climate change winners and losers highlights the rapid loss of cold‐dwelling species. Diversity Distributions. 2016;22(4):468–80. https://doi.org/10.1111/ddi.12412 64. Prieto-Torres DA, Sánchez-González LA, Ortiz-Ramírez MF, Ramírez-Albores JE, García-Trejo EA, Navarro-Sigüenza AG. Climate warming affects spatio-temporal biodiversity patterns of a highly vulnerable Neotropical avifauna. Climatic Change. 2021;165(3–4). https://doi.org/10.1007/ s10584-021-03091-3 65. Palmer G, Platts PJ, Brereton T, Chapman JW, Dytham C, Fox R, et al. Climate change, climatic variation and extreme biological responses. Philos Trans R Soc Lond B Biol Sci. 2017;372(1723):20160144. https://doi.org/10.1098/rstb.2016.0144 PMID: 28483874

PLOS One | https://doi.org/10.1371/journal.pone.0330153 August 29, 2025 19 / 20 66. Habel JC, Ulrich W, Gros P, Teucher M, Schmitt T. Butterfly species respond differently to climate warming and land use change in the northern Alps. Sci Total Environ. 2023;890:164268. https://doi.org/10.1016/j.scitotenv.2023.164268 PMID: 37211129 67. Devictor V, Julliard R, Couvet D, Jiguet F. Birds are tracking climate warming, but not fast enough. Proc Biol Sci. 2008;275(1652):2743–8. https:// doi.org/10.1098/rspb.2008.0878 PMID: 18713715 68. Rumpf SB, Hülber K, Zimmermann NE, Dullinger S. Elevational rear edges shifted at least as much as leading edges over the last century. Global Ecol Biogeogr. 2018;28(4):533–43. https://doi.org/10.1111/geb.12865 69. Harsch MA, HilleRisLambers J. Climate warming and seasonal precipitation change interact to limit species distribution shifts across Western North America. PLoS One. 2016;11(7):e0159184. https://doi.org/10.1371/journal.pone.0159184 PMID: 27447834 70. Herrando S, Titeux N, Brotons L, Anton M, Ubach A, Villero D, et al. Contrasting impacts of precipitation on Mediterranean birds and butterflies. Sci Rep. 2019;9(1):5680. https://doi.org/10.1038/s41598-019-42171-4 PMID: 30952919 71. Tilman D, May RM, Lehman CL, Nowak MA. Habitat destruction and the extinction debt. Nature. 1994;371(6492):65–6. https://doi. org/10.1038/371065a0 72. Barlow J, Lennox GD, Ferreira J, Berenguer E, Lees AC, Mac Nally R, et al. Anthropogenic disturbance in tropical forests can double biodiversity

loss from deforestation. Nature. 2016;535(7610):144–7. https://doi.org/10.1038/nature18326 PMID: 27362236 73. Betts MG, Wolf C, Ripple WJ, Phalan B, Millers KA, Duarte A, et al. Global forest loss disproportionately erodes biodiversity in intact landscapes. Nature. 2017;547(7664):441–4. https://doi.org/10.1038/nature23285 PMID: 28723892 74. Davey CM, Devictor V, Jonzén N, Lindström A, Smith HG. Impact of climate change on communities: revealing species’ contribution. J Anim Ecol. 2013;82(3):551–61. https://doi.org/10.1111/1365-2656.12035 PMID: 23398634 75. Tscharntke T, Sekercioglu CH, Dietsch TV, Sodhi NS, Hoehn P, Tylianakis JM. Landscape constraints on functional diversity of birds and insects in tropical agroecosystems. Ecology. 2008;89(4):944–51. https://doi.org/10.1890/07-0455.1 PMID: 18481519 76. Fontana V, Guariento E, Hilpold A, Niedrist G, Steinwandter M, Spitale D, et al. Species richness and beta diversity patterns of multiple taxa along an elevational gradient in pastured grasslands in the European Alps. Sci Rep. 2020;10(1):12516. https://doi.org/10.1038/s41598-020-69569-9 PMID: 32719437 77. García‐Navas V, Sattler T, Schmid H, Ozgul A. Temporal homogenization of functional and beta diversity in bird communities of the Swiss Alps. Diversity Distributions. 2020;26(8):900–11. https://doi.org/10.1111/ddi.13076 78. Si X, Baselga A, Ding P. Revealing Beta-diversity patterns of breeding bird and lizard communities on inundated land-bridge islands by

separating the turnover and nestedness components. PLoS One. 2015;10(5):e0127692. https://doi.org/10.1371/journal.pone.0127692 PMID: 25992559 79. Soininen J, Heino J, Wang J. A meta‐analysis of nestedness and turnover components of beta diversity across organisms and ecosystems. Global Ecol Biogeogr. 2017;27(1):96–109. https://doi.org/10.1111/geb.12660 80. Staude IR, Vélez‐Martin E, Andrade BO, Podgaiski LR, Boldrini II, Mendonça M Jr, et al. Local biodiversity erosion in south Brazilian grasslands under moderate levels of landscape habitat loss. J Appl Ecol. 2018;55(3):1241–51. https://doi.org/10.1111/1365-2664.13067 81. Freeman BG, Scholer MN, Ruiz-Gutierrez V, Fitzpatrick JW. Climate change causes upslope shifts and mountaintop extirpations in a tropical bird community. Proc Natl Acad Sci U S A. 2018;115(47):11982–7. https://doi.org/10.1073/pnas.1804224115 PMID: 30373825 82. Bowler D, Richter RL, Eskildsen D, Kamp J, Moshøj CM, Reif J, et al. Geographic variation in the population trends of common breeding birds across central Europe. Basic Appl Ecol. 2021;56:72–84. https://doi.org/10.1016/j.baae.2021.07.004 83. McDonald KW, McClure CJW, Rolek BW, Hill GE. Diversity of birds in eastern North America shifts north with global warming. Ecol Evol. 2012;2(12):3052–60. https://doi.org/10.1002/ece3.410 PMID: 23301172 84. Princé K, Zuckerberg B. Climate change in our backyards: the reshuffling of North America’s winter bird communities. Glob Chang Biol. 2015;21(2):572–85. https://doi.org/10.1111/gcb.12740 PMID: 25322929

85. Hillebrand H, Blasius B, Borer ET, Chase JM, Downing JA, Eriksson BK, et al. Biodiversity change is uncoupled from species richness trends: consequences for conservation and monitoring. J Appl Ecol. 2017;55(1):169–84. https://doi.org/10.1111/1365-2664.12959 86. Concepción ED, Obrist MK, Moretti M, Altermatt F, Baur B, Nobis MP. Impacts of urban sprawl on species richness of plants, butterflies, gastropods and birds: not only built-up area matters. Urban Ecosyst. 2015;19(1):225–42. https://doi.org/10.1007/s11252-015-0474-4 87. Devictor V, Julliard R, Couvet D, Lee A, Jiguet F. Functional homogenization effect of urbanization on bird communities. Conserv Biol. 2007;21(3):741–51. https://doi.org/10.1111/j.1523-1739.2007.00671.x PMID: 17531052 88. Mimet A, Buitenwerf R, Sandel B, Svenning J, Normand S. Recent global changes have decoupled species richness from specialization patterns in North American birds. Global Ecol Biogeogr. 2019;28(11):1621–35. https://doi.org/10.1111/geb.12978 89. Rabbetts M, Fahrig L, Mitchell GW, Hannah KC, Collins SJ, Wilson S. Direct and indirect effects of agricultural land cover on avian biodiversity in eastern Canada. Biodivers Conserv. 2023;32(4):1403–21. https://doi.org/10.1007/s10531-023-02559-1 PMID: 36992920 90. Rigal S, Dakos V, Alonso H, Auniņš A, Benkő Z, Brotons L, et al. Farmland practices are driving bird population decline across Europe. Proc Natl Acad Sci U S A. 2023;120(21):e2216573120. https://doi.org/10.1073/pnas.2216573120 PMID: 37186854

PLOS One | https://doi.org/10.1371/journal.pone.0330153 August 29, 2025 20 / 20 91. Dornelas M, Gotelli NJ, McGill B, Shimadzu H, Moyes F, Sievers C, et al. Assemblage time series reveal biodiversity change but not systematic loss. Science. 2014;344(6181):296–9. https://doi.org/10.1126/science.1248484 PMID: 24744374 92. Vellend M, Baeten L, Myers-Smith IH, Elmendorf SC, Beauséjour R, Brown CD, et al. Global meta-analysis reveals no net change in local-scale plant biodiversity over time. Proc Natl Acad Sci U S A. 2013;110(48):19456–9. https://doi.org/10.1073/pnas.1312779110 PMID: 24167259 93. Northrup JM, Rivers JW, Yang Z, Betts MG. Synergistic effects of climate and land-use change influence broad-scale avian population declines. Glob Chang Biol. 2019;25(5):1561–75. https://doi.org/10.1111/gcb.14571 PMID: 30810257