Philippa Kaur

1.Simpson, G. G. History of the fauna of Latin America. Am. Sci. 38, 361–389 (1950).
Google Scholar 
2.Patterson, B. D. & Costa, L. P. Bones, Clones, and Biomes: The History and Geography of Recent Neotropical Mammals (The University of Chicago Press, 2012).3.Cidade, G. M., Fortier, D. & Hsiou, A. S. The crocodylomorph fauna of the Cenozoic of South America and its evolutionary history: a review. J. South Am. Earth Sci. 90, 392–411 (2019).ADS 

Google Scholar 
4.Tambussi, C. P. & Degrange, F. J. South American and Antarctic Continental Cenozoic Birds. (Springer Netherlands, 2013). https://doi.org/10.1007/978-94-007-5467-6.5.Marshall, L. G. Evolution of the carnivorous adaptive zone in South America. In Major Patterns in Vertebrate Evolution (eds. Hecht, M. K., Goody, P. C. & Hecht, B. M.) 709–721 (Springer US, 1977). https://doi.org/10.1007/978-1-4684-8851-7_24.6.Marshall, L. G. & Cifelli, R. L. Analysis of changing diversity patterns in Cenozoic land mammal age faunas, South America. Palaeovertebrata 19, 169–210 (1990).
Google Scholar 
7.Prevosti, F. J., Forasiepi, A. & Zimicz, N. The evolution of the cenozoic terrestrial mammalian predator guild in South America: competition or replacement?. J. Mamm. Evol. 20, 3–21 (2013).
Google Scholar 
8.Prevosti, F. J. & Forasiepi, A. M. Evolution of South American mammalian predators during the Cenozoic: paleobiogeographic and paleoenvironmental contingencies. (Springer International Publishing, 2018). https://doi.org/10.1007/978-3-319-03701-1.9.Croft, D. A. Do marsupials make good predators? Insights from predator-prey diversity ratios. Evol. Ecol. Res. 8, 1193–1214 (2006).
Google Scholar 
10.Albino, A. M. Snakes from the Paleocene and Eocene of Patagonia (Argentina): Paleoecology and coevolution with mammals. Hist. Biol. 7, 51–69 (1993).
Google Scholar 
11.Head, J. J. et al. Giant boid snake from the Palaeocene neotropics reveals hotter past equatorial temperatures. Nature 457, 715–717 (2009).ADS 
CAS 
PubMed 

Google Scholar 
12.de Muizon, C., Ladevèze, S., Selva, C., Vignaud, R. & Goussard, F. Allqokirus australis (Sparassodonta, Metatheria) from the early Palaeocene of Tiupampa (Bolivia) and the rise of the metatherian carnivorous radiation in South America. Geodiversitas 40, 363 (2018).
Google Scholar 
13.Forasiepi, A. M. Osteology of Arctodictis sinclairi (Mammalia, Metatheria, Sparassodonta) and phylogeny of Cenozoic metatherian carnivores from South America. Monogr. Mus. Argentino Cienc. Nat., n.s. 6, 1–174 (2009).14.Prevosti, F. J. et al. New radiometric 40Ar–39Ar dates and faunistic analyses refine evolutionary dynamics of Neogene vertebrate assemblages in southern South America. Sci. Rep. 11, 9830 (2021).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
15.Pérez, L. F. et al. Oceanographic and climatic consequences of the tectonic evolution of the southern scotia sea basins, Antarctica. Earth-Science Rev. 198, 102922 (2019).
Google Scholar 
16.Williams, S. E., Whittaker, J. M., Halpin, J. A. & Müller, R. D. Australian-Antarctic breakup and seafloor spreading: Balancing geological and geophysical constraints. Earth-Science Rev. 188, 41–58 (2019).ADS 

Google Scholar 
17.Gelfo, J. N. Considerations about the evolutionary stasis of Notiolofos arquinotiensis (Mammalia: Sparnotheriodontidae), Eocene of Seymour Island, Antartica. Ameghiniana 53, 316–332 (2016).
Google Scholar 
18.Goin, F. J. et al. New metatherian mammal from the Early Eocene of Antarctica. J. Mamm. Evol. https://doi.org/10.1007/s10914-018-9449-6 (2018).Article 

Google Scholar 
19.Reguero, M. A. et al. Final Gondwana breakup: The Paleogene South American native ungulates and the demise of the South America-Antarctica land connection. Glob. Planet. Change 123, 400–413 (2014).ADS 

Google Scholar 
20.Ramos, V. A. & Aleman, A. Tectonic evolution of the Andes. in Tectonic evolution of South America (eds. Cordani, U. G., Milani, E. J., Thomaz Filho, A. & Campos, D. A.) 635–685 (2000).21.Boschman, L. M. Andean mountain building since the Late Cretaceous: A paleoelevation reconstruction. Earth-Science Rev. 220, 103640 (2021).
Google Scholar 
22.Fosdick, J. C., Reat, E. J., Carrapa, B., Ortiz, G. & Alvarado, P. M. Retroarc basin reorganization and aridification during Paleogene uplift of the southern central Andes. Tectonics 36, 493–514 (2017).ADS 

Google Scholar 
23.Leier, A., McQuarrie, N., Garzione, C. & Eiler, J. Stable isotope evidence for multiple pulses of rapid surface uplift in the Central Andes, Bolivia. Earth Planet. Sci. Lett. 371–372, 49–58 (2013).ADS 

Google Scholar 
24.Cione, A. L., Gasparini, G. M., Soibelzon, E., Soibelzon, L. H. & Tonni, E. P. The Great American Biotic Interchange: a South American perspective. (Springer, 2015).25.Coates, A. G. & Stallard, R. F. How old is the isthmus of Panama?. Bull. Mar. Sci. 89, 801–813 (2013).
Google Scholar 
26.Montes, C. et al. Middle Miocene closure of the Central American Seaway. Science (80-) 348, 226–229 (2015).ADS 
CAS 

Google Scholar 
27.Auderset, A. et al. Gulf Stream intensification after the early Pliocene shoaling of the Central American Seaway. Earth Planet. Sci. Lett. 520, 268–278 (2019).CAS 

Google Scholar 
28.Scher, H. D. et al. Onset of Antarctic Circumpolar Current 30 million years ago as Tasmanian Gateway aligned with westerlies. Nature 523, 580–583 (2015).ADS 
CAS 
PubMed 

Google Scholar 
29.Benton, M. J. The Red Queen and the Court Jester: species diversity and the role of biotic and abiotic factors through time. Science (80-) 323, 728–732 (2009).ADS 
CAS 

Google Scholar 
30.Van Valen, L. New evolutionary law. Evol. Theory 1, 1–30 (1973).
Google Scholar 
31.Cione, A. L., Tonni, E. P. & Soibelzon, L. The Broken Zig-Zag: Late Cenozoic large mammal and tortoise extinction in South America. Rev. del Mus. Argentino Ciencias Nat. n.s. 5, 1–19 (2003).32.Leigh, E. G., O’Dea, A. & Vermeij, G. J. Historical biogeography of the isthmus of Panama. Biol. Rev. 89, 148–172 (2014).PubMed 

Google Scholar 
33.Werdelin, L. Jaw geometry and molar morphology in Marsupial carnivores: Analysis of a constraint and its macroevolutionary consequences. Paleobiology 13, 342–350 (1987).
Google Scholar 
34.Goin, F. J. & Montalvo, C. Revisión sistemática y reconocimiento de una nueva especie del género Thylatheridium Reig (Marsupialia, Didelphidae). Ameghiniana 25, 161–167 (1988).
Google Scholar 
35.Goin, F. J. & Pardiñas, U. F. J. Revision de las especies del genero Hyperdidelphys Ameghino, 1904 (Mammalia, Marsupialia, Didelphidae). Su significación filogenética, estratigráfica y adaptativa en el Neogeno del Cono Sur Sudamericano. Estud. Geológicos 52, 327–359 (1996).36.Beck, R. M. D. & Taglioretti, M. L. A nearly complete juvenile skull of the marsupial Sparassocynus derivatus from the Pliocene of Argentina, the affinities of “sparassocynids”, and the diversification of opossums (Marsupialia; Didelphimorphia; Didelphidae). J. Mamm. Evol. 27, 385–417 (2020).
Google Scholar 
37.López-Aguirre, C., Archer, M., Hand, S. J. & Laffan, S. W. Extinction of South American sparassodontans (Metatheria): Environmental fluctuations or complex ecological processes?. Palaeontology 60, 91–115 (2017).
Google Scholar 
38.Forasiepi, A. M., Martinelli, A. G. & Goin, F. J. Revisión taxonómica de Parahyaenodon argentinus Ameghino y sus implicancias en el conocimiento de los grandes mamíferos carnívoros del Mio-Plioceno de América de Sur. Ameghiniana 44, 143–159 (2007).
Google Scholar 
39.Zimicz, N. Avoiding competition: the ecological history of late Cenozoic metatherian carnivores in South America. J. Mamm. Evol. 21, 383–393 (2014).
Google Scholar 
40.Echarri, S., Ercoli, M. D., Chemisquy, M. A., Turazzini, G. & Prevosti, F. J. Mandible morphology and diet of the South American extinct metatherian predators (Mammalia, Metatheria, Sparassodonta). Earth Environ. Sci. Trans. R. Soc. Edinburgh 106, 277–288 (2017).41.Croft, D. A., Engelman, R. K., Dolgushina, T. & Wesley, G. Diversity and disparity of sparassodonts (Metatheria) reveal non-analogue nature of ancient South American mammalian carnivore guilds. Proc. R. Soc. B Biol. Sci. 285, 20172012 (2018).
Google Scholar 
42.Engelman, R. K. & Croft, D. A. A new species of small-bodied sparassodont (Mammalia, Metatheria) from the middle Miocene locality of Quebrada Honda, Bolivia. J. Vertebr. Paleontol. 34, 672–688 (2014).
Google Scholar 
43.Engelman, R. K. & Croft, D. A. Strangers in a strange land: ecological dissimilarity to metatherian carnivores may partly explain early colonization of South America by Cyonasua- group procyonids. Paleobiology 45, 598–611 (2019).
Google Scholar 
44.Engelman, R. K., Anaya, F. & Croft, D. A. Australogale leptognathus, gen. et sp. nov., a Second Species of Small Sparassodont (Mammalia: Metatheria) from the Middle Miocene Locality of Quebrada Honda, Bolivia. J. Mamm. Evol. 27, 37–54 (2020).45.Silvestro, D., Schnitzler, J., Liow, L. H., Antonelli, A. & Salamin, N. Bayesian estimation of speciation and extinction from incomplete fossil occurrence data. Syst. Biol. 63, 349–367 (2014).PubMed 

Google Scholar 
46.Lehtonen, S. et al. Environmentally driven extinction and opportunistic origination explain fern diversification patterns. Sci. Rep. 7, 4831 (2017).ADS 
PubMed 
PubMed Central 

Google Scholar 
47.Rangel, C. et al. Diversity, affinities and adaptations of the basal sparassodont Patene Simpson, 1935 (Mammalia, Metatheria). Ameghiniana 56, 263–289 (2019).
Google Scholar 
48.Alroy, J. et al. Effects of sampling standardization on estimates of Phanerozoic marine diversification. Proc. Natl. Acad. Sci. 98, 6261–6266 (2001).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
49.Badgley, C. The multiple scales of biodiversity. Paleobiology 29, 11–13 (2003).
Google Scholar 
50.Behrensmeyer, A. K., Kidwell, S. M. & Gastaldo, R. A. Taphonomy and paleobiology. Paleobiology 26, 103–147 (2000).
Google Scholar 
51.Butler, R. J., Barrett, P. M., Nowbath, S. & Upchurch, P. Estimating the effects of sampling biases on pterosaur diversity patterns: implications for hypotheses of bird/pterosaur competitive replacement. Paleobiology 35, 432–446 (2009).
Google Scholar 
52.Crampton, J. S. et al. Estimating the rock volume bias in paleobiodiversity studies. Science (80-) 301, 358–360 (2003).ADS 
CAS 

Google Scholar 
53.Kalmar, A. & Currie, D. J. The completeness of the continental fossil record and its impact on patterns of diversification. Paleobiology 36, 51–60 (2010).
Google Scholar 
54.Newham, E., Benson, R., Upchurch, P. & Goswami, A. Mesozoic mammaliaform diversity: the effect of sampling corrections on reconstructions of evolutionary dynamics. Palaeogeogr. Palaeoclimatol. Palaeoecol. 412, 32–44 (2014).
Google Scholar 
55.Prevosti, F. J. & Soibelzon, L. H. Evolution of the South American carnivores (Mammalia, Carnivora): a paleontological perspective. In Bones, Clones, and Biomes: The History and Geography of Recent Neotropical Mammals (eds. Patterson, B. D. & Costa, L. P.) 102–122 (University of Chicago Press, 2012).56.Carrillo, J. D., Forasiepi, A., Jaramillo, C. & Sánchez-Villagra, M. R. Neotropical mammal diversity and the Great American Biotic Interchange: spatial and temporal variation in South America’s fossil record. Front. Genet. 5, 451 (2015).PubMed 
PubMed Central 

Google Scholar 
57.Silvestro, D., Salamin, N., Antonelli, A. & Meyer, X. Improved estimation of macroevolutionary rates from fossil data using a Bayesian framework. Paleobiology https://doi.org/10.1017/pab.2019.23 (2019).Article 

Google Scholar 
58.Condamine, F. L., Guinot, G., Benton, M. J. & Currie, P. J. Dinosaur biodiversity declined well before the asteroid impact, influenced by ecological and environmental pressures. Nat. Commun. 12, 3833 (2021).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
59.Silvestro, D., Antonelli, A., Salamin, N. & Quental, T. B. The role of clade competition in the diversification of North American canids. Proc. Natl. Acad. Sci. 112, 8684–8689 (2015).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
60.Alroy, J. Constant extinction, constrained diversification, and uncoordinated stasis in North American mammals. Palaeogeogr. Palaeoclimatol. Palaeoecol. 127, 285–311 (1996).
Google Scholar 
61.Moen, D. & Morlon, H. Why does diversification slow down?. Trends Ecol. Evol. 29, 190–197 (2014).PubMed 

Google Scholar 
62.Bromham, L. The genome as a life-history character: why rate of molecular evolution varies between mammal species. Philos. Trans. R. Soc. B Biol. Sci. 366, 2503–2513 (2011).
Google Scholar 
63.Feng, P. & Zhou, Q. Absence of relationship between mitochondrial DNA evolutionary rate and longevity in mammals except for CYTB. J. Mamm. Evol. 26, 1–7 (2019).
Google Scholar 
64.Gillooly, J. F., Allen, A. P., West, G. B. & Brown, J. H. The rate of DNA evolution: Effects of body size and temperature on the molecular clock. Proc. Natl. Acad. Sci. 102, 140–145 (2005).ADS 
CAS 
PubMed 

Google Scholar 
65.Martin, A. P. & Palumbi, S. R. Body size, metabolic rate, generation time, and the molecular clock. Proc. Natl. Acad. Sci. 90, 4087–4091 (1993).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
66.Barraclough, T. G., Vogler, A. P. & Harvey, P. H. Revealing the factors that promote speciation. Philos. Trans. R. Soc. Lond. Ser. B Biol. Sci. 353, 241–249 (1998).67.Barraclough, T. G. & Savolainen, V. Evolutionary rates and species diversity in flowering plants. Evolution (N. Y) 55, 677–683 (2001).CAS 

Google Scholar 
68.Raia, P., Passaro, F., Fulgione, D. & Carotenuto, F. Habitat tracking, stasis and survival in Neogene large mammals. Biol. Lett. 8, 64–66 (2012).CAS 
PubMed 

Google Scholar 
69.Viranta, S. Geographic and temporal ranges of middle and late Miocene Carnivores. J. Mammal. 84, 1267–1278 (2003).
Google Scholar 
70.Flynn, L. J. et al. Neogene Siwalik mammalian lineages: Species longevities, rates of change, and modes of speciation. Palaeogeogr. Palaeoclimatol. Palaeoecol. 115, 249–264 (1995).
Google Scholar 
71.Van Valen, L. Group selection, sex, and fossils. Evolution (N. Y) 29, 87 (1975).
Google Scholar 
72.Cardillo, M. Biological determinants of extinction risk: why are smaller species less vulnerable?. Anim. Conserv. 6, 63–69 (2003).
Google Scholar 
73.Liow, L. H. et al. Higher origination and extinction rates in larger mammals. Proc. Natl. Acad. Sci. 105, 6097–6102 (2008).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
74.McLain, D. K. Cope’s rules, sexual selection, and the loss of ecological plasticity. Oikos 68, 490–500 (1993).
Google Scholar 
75.Hautmann, M. What is macroevolution?. Palaeontology 63, 1–11 (2020).
Google Scholar 
76.Stanley, S. M. The general correlation between rate of speciation and rate of extinction: fortuitous causal linkages. In Causes of evolution: A paleontological perspective (eds. Ross, R. M. & Allmon, W. D.) 103–127 (University of Chicago Press, 1990).77.Marshall, C. R. Five palaeobiological laws needed to understand the evolution of the living biota. Nat. Ecol. Evol. 1, 0165 (2017).
Google Scholar 
78.Ercoli, M. D., Prevosti, F. J. & Forasiepi, A. M. The structure of the mammalian predator guild in the Santa Cruz Formation (late Early Miocene). J. Mamm. Evol. 21, 369–381 (2014).
Google Scholar 
79.Marshall, L. G. Evolution of the Borhyaenidae, extinct south american predaceous marsupials. Univ. Calif. Publ. Geol. Sci. 17, 1–89 (1978).
Google Scholar 
80.Prevosti, F. J., Forasiepi, A. M., Ercoli, M. D. & Turazzini, G. F. Paleoecology of the mammalian carnivores (Metatheria, Sparassodonta) of the Santa Cruz Formation (late Early Miocene). In Early Miocene paleobiology in Patagonia (eds. Vizcaino, S. F., Kay, R. F. & Bargo, M. S.) 173–193 (Cambridge University Press, 2012). https://doi.org/10.1017/CBO9780511667381.012.81.Carbone, C., Teacher, A. & Rowcliffe, J. M. The costs of carnivory. PLoS Biol. 5, e22 (2007).PubMed 
PubMed Central 

Google Scholar 
82.Carbone, C., Mace, G. M., Roberts, S. C. & Macdonald, D. W. Energetic constraints on the diet of terrestrial carnivores. Nature 402, 286–288 (1999).ADS 
CAS 
PubMed 

Google Scholar 
83.Rovinsky, D. S., Evans, A. R., Martin, D. G. & Adams, J. W. Did the thylacine violate the costs of carnivory? Body mass and sexual dimorphism of an iconic Australian marsupial. Proc. R. Soc. B Biol. Sci. 287, 20201537 (2020).
Google Scholar 
84.Van Valkenburgh, B., Wang, X. & Damuth, J. Cope’s Rule, hypercarnivory, and extinction in North American canids. Science (80-) 306, 101–104 (2004).ADS 

Google Scholar 
85.Finarelli, J. A. Mechanisms behind active trends in body size evolution of the Canidae (Carnivora: Mammalia). Am. Nat. 170, 876–885 (2007).PubMed 

Google Scholar 
86.Cione, A. L. & Tonni, E. P. Chronostratigraphy and “Land-Mammal Ages” in the Cenozoic of southern South America: principles, practices, and the “Uquian” problem. J. Paleontol. 69, 135–159 (1995).
Google Scholar 
87.Soibelzon, L. H. & Prevosti, F. J. Los carnívoros (Carnivora, Mammalia) terrestres del Cuaternario de América del Sur. in Geomorfología Litoral i Quaternari. Homenatge a Joan Cuerda Barceló. Mon. Soc. Hist. Nat. (eds. Pons, G. X. & Vicens, D.) vol. 14 49–68 (2007).88.Argot, C. Evolution of South American mammalian predators (Borhyaenoidea): Anatomical and palaeobiological implications. Zool. J. Linn. Soc. 140, 487–521 (2004).
Google Scholar 
89.Degrange, F. J. Hind limb morphometry of terror birds (Aves, Cariamiformes, Phorusrhacidae): functional implications for substrate preferences and locomotor lifestyle. Earth Environ. Sci. Trans. R. Soc. Edinburgh 106, 257–276 (2015).90.Angst, D., Buffetaut, E., Lecuyer, C. & Amiot, R. A new method for estimating locomotion type in large ground birds. Palaeontology 59, 217–223 (2016).
Google Scholar 
91.Gurevitch, J. & Padilla, D. K. Are invasive species a major cause of extinctions?. Trends Ecol. Evol. 19, 470–474 (2004).PubMed 

Google Scholar 
92.Davis, M. A. Biotic globalization: Does competition from introduced species threaten biodiversity?. Bioscience 53, 481–489 (2003).
Google Scholar 
93.Prowse, T. A. A., Johnson, C. N., Bradshaw, C. J. A. & Brook, B. W. An ecological regime shift resulting from disrupted predator–prey interactions in Holocene Australia. Ecology 95, 693–702 (2014).PubMed 

Google Scholar 
94.Marshall, L. G. A new species of Lycopsis (Borhyaenidae: Marsupialia) from the La Venta Fauna (Late Miocene) of Colombia, South America. J. Paleontol. 51, 633–642 (1977).
Google Scholar 
95.Tomassini, R. L., Montalvo, C. I., Bargo, M. S., Vizcaíno, S. F. & Cuitiño, J. I. Sparassodonta (Metatheria) coprolites from the early-mid Miocene (Santacrucian age) of Patagonia (Argentina) with evidence of exploitation by coprophagous insects. Palaios 34, 639–651 (2019).ADS 

Google Scholar 
96.Bond, M., Cerdeño, E. & López, G. Los ungulados nativos de América del Sur. in Evolución biológica y climática de la región Pampeana durante los últimos cinco millones de años (eds. Alberdi, M. T., Leone, G. & Tonni, E. P.) 258–275 (Museo Nacional de Ciencias Naturales, 1995).97.Croft, D. A., Gelfo, J. N. & López, G. M. Splendid innovation: The extinct South American native ungulates. Annu. Rev. Earth Planet. Sci. 48, 259–290 (2020).ADS 
CAS 

Google Scholar 
98.Pascual, R. & Jaureguizar, E. O. Evolving climates and mammal faunas in cenozoic South America. J. Hum. Evol. 19, 23–60 (1990).
Google Scholar 
99.Patterson, B. & Pascual, R. The fossil mammal fauna of South America. Q. Rev. Biol. 43, 409–451 (1968).
Google Scholar 
100.Miller, K. G. et al. Cenozoic sea-level and cryospheric evolution from deep-sea geochemical and continental margin records. Sci. Adv. 6, eaaz1346 (2020).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
101.Rae, J. W. B. et al. Atmospheric CO2 over the past 66 million years from marine archives. Annu. Rev. Earth Planet. Sci. 49, 609–641 (2021).ADS 
CAS 

Google Scholar 
102.Insel, N., Poulsen, C. J. & Ehlers, T. A. Influence of the Andes mountains on South American moisture transport, convection, and precipitation. Clim. Dyn. 35, 1477–1492 (2010).
Google Scholar 
103.Garzione, C. N. et al. Tectonic evolution of the Central Andean Plateau and implications for the growth of plateaus. Annu. Rev. Earth Planet. Sci. 45, 529–559 (2017).ADS 
CAS 

Google Scholar 
104.Lossada, A. C. et al. Cenozoic uplift and exhumation of the Frontal Cordillera between 30° and 35° S and the influence of the subduction dynamics in the flat slab subduction context, South Central Andes. In The Evolution of the Chilean-Argentinean Andes (eds. Folguera, A. et al.) 387–409 (Springer Earth System Sciences, 2018). https://doi.org/10.1007/978-3-319-67774-3_16.105.Mora, A. et al. Tectonic history of the Andes and sub-Andean zones: implications for the development of the Amazon drainage basin. In Amazonia: Landscape and Species Evolution (eds. Hoorn, C. & Wesselingh, F. P.) 38–60 (Wiley-Blackwell Publishing Ltd., 2011). https://doi.org/10.1002/9781444306408.ch4.106.Pingel, H. et al. Late Cenozoic topographic evolution of the Eastern Cordillera and Puna Plateau margin in the southern Central Andes (NW Argentina). Earth Planet. Sci. Lett. 535, 116112 (2020).CAS 

Google Scholar 
107.Takahashi, K. & Battisti, D. S. Processes controlling the mean tropical Pacific precipitation pattern. Part I: The Andes and the Eastern Pacific ITCZ. J. Clim. 20, 3434–3451 (2007).ADS 

Google Scholar 
108.Amidon, W. H. et al. Mio-Pliocene aridity in the south-central Andes associated with Southern Hemisphere cold periods. Proc. Natl. Acad. Sci. 114, 6474–6479 (2017).CAS 
PubMed 
PubMed Central 

Google Scholar 
109.Carrapa, B., Clementz, M. & Feng, R. Ecological and hydroclimate responses to strengthening of the Hadley circulation in South America during the Late Miocene cooling. Proc. Natl. Acad. Sci. 116, 9747–9752 (2019).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
110.Hartley, A. J. Andean uplift and climate change. J. Geol. Soc. Lond. 160, 7–10 (2003).
Google Scholar 
111.Barreda, V., Guler, V. & Palazzesi, L. Late Miocene continental and marine palynological assemblages from Patagonia. Dev. Quat. Sci. 11, 343–350 (2008).
Google Scholar 
112.Barreda, V. & Palazzesi, L. Patagonian vegetation turnovers during the Paleogene-early Neogene: Origin of arid-adapted floras. Bot. Rev. 73, 31–50 (2007).
Google Scholar 
113.Ortiz-Jaureguizar, E. & Cladera, G. A. Paleoenvironmental evolution of southern South America during the Cenozoic. J. Arid Environ. 66, 498–532 (2006).ADS 

Google Scholar 
114.Krockenberger, A. Lactation. In Marsupials (eds. Armati, P. J., Dickman, C. R. & Hume, I. D.) 108–136 (Cambridge University Press, 2006). https://doi.org/10.1017/CBO9780511541889.005.115.Morton, S. R., Recher, H. F., Thompson, S. D. & Braithwaite, R. W. Comments on the relative advantages of marsupial and eutherian reproduction. Am. Nat. 120, 128–134 (1982).
Google Scholar 
116.Thompson, S. D. Body size, duration of parental care, and the intrinsic rate of natural increase in eutherian and metatherian mammals. Oecologia 71, 201–209 (1987).ADS 
CAS 
PubMed 

Google Scholar 
117.Holloway, J. C. & Geiser, F. Seasonal changes in the thermoenergetics of the marsupial sugar glider Petaurus breviceps. J. Comp. Physiol. B Biochem. Syst. Environ. Physiol. 171, 643–650 (2001).CAS 

Google Scholar 
118.Sánchez-Villagra, M. R. Why are there fewer marsupials than placentals? On the relevance of geography and physiology to evolutionary patterns of mammalian diversity and disparity. J. Mamm. Evol. 20, 279–290 (2013).
Google Scholar 
119.Bennett, C. V., Upchurch, P., Goin, F. J. & Goswami, A. Deep time diversity of metatherian mammals: Implications for evolutionary history and fossil-record quality. Paleobiology 44, 171–198 (2018).
Google Scholar 
120.Goin, F. J., Woodburne, M. O., Zimicz, A. N., Martin, G. M. & Chornogubsky, L. A Brief History of South American Metatherians: Evolutionary Contexts and Intercontinental Dispersals (Springer, 2016).121.Jablonski, D. Biotic interactions and macroevolution: Extensions and mismatches across scales and levels. Evolution (N. Y) 62, 715–739 (2008).
Google Scholar 
122.Mills, B. J. W., Belcher, C. M., Lenton, T. M. & Newton, R. J. A modeling case for high atmospheric oxygen concentrations during the Mesozoic and Cenozoic. Geology 44, 1023–1026 (2016).ADS 
CAS 

Google Scholar 
123.Westerhold, T. et al. An astronomically dated record of Earth’s climate and its predictability over the last 66 million years. Science (80-) 369, 1383–1387 (2020).ADS 
CAS 

Google Scholar 
124.Silvestro, D., Salamin, N. & Schnitzler, J. PyRate: A new program to estimate speciation and extinction rates from incomplete fossil data. Methods Ecol. Evol. 5, 1126–1131 (2014).
Google Scholar 
125.Green, P. J. Reversible jump Markov chain Monte Carlo computation and Bayesian model determination. Biometrika 82, 711–732 (1995).MathSciNet 
MATH 

Google Scholar 
126.Rambaut, A., Drummond, A. J., Xie, D., Baele, G. & Suchard, M. A. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Syst. Biol. 67, 901–904 (2018).CAS 
PubMed 
PubMed Central 

Google Scholar  More

1.Mesoudi, A. & Thornton, A. What is cumulative cultural evolution? Proc. R. Soc. B 285, 20180712 (2018).PubMed 
PubMed Central 

Google Scholar 
2.Schofield, D. P., McGrew, W. C., Takahashi, A. & Hirata, S. Cumulative culture in nonhumans: overlooked findings from Japanese monkeys? Primates 59, 113–122 (2018).PubMed 

Google Scholar 
3.Boesch, C. & Tomasello, M. Chimpanzee and human cultures. Curr. Anthropol. 39, 591–614 (1998).
Google Scholar 
4.Tennie, C., Call, J. & Tomasello, M. Ratcheting up the ratchet: on the evolution of cumulative culture. Philos. Trans. R. Soc. B 364, 2405–2415 (2009).
Google Scholar 
5.Dean, L. G., Vale, G. L., Laland, K. N., Flynn, E. & Kendal, R. L. Human cumulative culture: a comparative perspective. Biol. Rev. Camb. Philos. Soc. 89, 284–301 (2014).PubMed 

Google Scholar 
6.Whiten, A., McGuigan, N., Marshall-Pescini, S. & Hopper, L. M. Emulation, imitation, over-imitation and the scope of culture for child and chimpanzee. Philos. Trans. R. Soc. B 364, 2417–2428 (2009).
Google Scholar 
7.Tennie, C., Bandini, E., van Schaik, C. P. & Hopper, L. M. The zone of latent solutions and its relevance to understanding ape cultures. Biol. Philos. 35, 55 (2020).PubMed 
PubMed Central 

Google Scholar 
8.Sasaki, T. & Biro, D. Cumulative culture can emerge from collective intelligence in animal groups. Nat. Commun. 8, 15049 (2017).CAS 
PubMed 
PubMed Central 

Google Scholar 
9.Musgrave, S. et al. Teaching varies with task complexity in wild chimpanzees. Proc. Natl Acad. Sci. USA 117, 969–976 (2020).CAS 
PubMed 

Google Scholar 
10.Boesch, C. et al. Chimpanzee ethnography reveals unexpected cultural diversity. Nat. Hum. Behav. 4, 910–916 (2020).PubMed 

Google Scholar 
11.Whiten, A. The burgeoning reach of animal culture. Science 372, eabe6514 (2021).CAS 
PubMed 

Google Scholar 
12.Whiten, A. et al. Cultures in chimpanzees. Nature 399, 682–685 (1999).CAS 

Google Scholar 
13.Sanz, C. & Morgan, D. Chimpanzee tool technology in the Goualougo Triangle, Republic of Congo. J. Hum. Evol. 52, 420–433 (2007).PubMed 

Google Scholar 
14.Whiten, A. & van Schaik, C. P. The evolution of animal ‘cultures’ and social intelligence. Philos. Trans. R. Soc. B 362, 603–620 (2007).
Google Scholar 
15.McGrew, W. C. The Cultured Chimpanzee: Reflections on Cultural Primatology (Cambridge Univ. Press, 2004).16.McGrew, W. C. Chimpanzee Material Culture: Implications for Human Evolution (Cambridge Univ. Press, 1992).17.Sanz, C., Schöning, C. & Morgan, D. Chimpanzees prey on army ants with specialized tool set. Am. J. Primatol. 72, 17–24 (2010).PubMed 

Google Scholar 
18.Reindl, E., Apperly, I. A., Beck, S. R. & Tennie, C. Young children copy cumulative technological design in the absence of action information. Sci. Rep. 7, 1788 (2017).CAS 
PubMed 
PubMed Central 

Google Scholar 
19.Neadle, D., Allritz, M. & Tennie, C. Food cleaning in gorillas: social learning is a possibility but not a necessity. PLoS ONE 12, e0188866 (2017).PubMed 
PubMed Central 

Google Scholar 
20.Tennie, C., Hopper, L. M. & van Schaik, C. P. in Chimpazees in Context (eds Hopper, L. M. & Ross, S. R.) 428–453 (Univ. of Chicago Press, 2020).21.Bandini, E. & Tennie, C. Spontaneous reoccurrence of “scooping”, a wild tool-use behaviour, in naive chimpanzees. PeerJ 5, e3814 (2017).PubMed 
PubMed Central 

Google Scholar 
22.Bandini, E. & Tennie, C. Individual acquisition of “stick pounding” behavior by naive chimpanzees. Am. J. Primatol. 81, e22987 (2019).PubMed 

Google Scholar 
23.Menzel, C., Fowler, A., Tennie, C. & Call, J. Leaf surface roughness elicits leaf swallowing behavior in captive chimpanzees (Pan troglodytes) and bonobos (P. paniscus), but not in gorillas (Gorilla gorilla) or orangutans (Pongo abelii). Int. J. Primatol. 34, 533–553 (2013).
Google Scholar 
24.Tonooka, R., Tomonaga, M. & Matsuzawa, T. Acquisition and transmission of tool making and use for drinking juice in a group of captive chimpanzees (Pan troglodytes). Jpn. Psychol. Res. 39, 253–265 (1997).
Google Scholar 
25.Celli, M. L., Hirata, S. & Tomonaga, M. Socioecological influences on tool use in captive chimpanzees. Int. J. Primatol. 25, 1267–1281 (2004).
Google Scholar 
26.Bernstein-Kurtycz, L. M., Hopper, L. M., Ross, S. R. & Tennie, C. Zoo-housed chimpanzees can spontaneously use tool sets but perseverate on previously successful tool-use methods. Anim. Behav. Cogn. 7, 288–309 (2020).
Google Scholar 
27.Boesch, C. in Evolution of Social Behaviour Patterns in Primates and Man (eds Runciman, W. G. et al.) 251–268 (Oxford Univ. Press, 1996).28.Neadle, D., Bandini, E. & Tennie, C. Testing the individual and social learning abilities of task-naïve captive chimpanzees (Pan troglodytes sp.) in a nut-cracking task. PeerJ 8, e8734 (2020).PubMed 
PubMed Central 

Google Scholar 
29.Biro, D. et al. Cultural innovation and transmission of tool use in wild chimpanzees: evidence from field experiments. Anim. Cogn. 6, 213–223 (2003).PubMed 

Google Scholar 
30.Boesch, C., Bombjaková, D., Meier, A. & Mundry, R. Learning curves and teaching when acquiring nut-cracking in humans and chimpanzees. Sci. Rep. 9, 1515 (2019).PubMed 
PubMed Central 

Google Scholar 
31.Matsuzawa, T. in Chimpanzee Cultures (eds Wrangham, R. W. et al.) 351–370 (Harvard Univ. Press, 1994).32.Matsuzawa, T. in Oxford Research Encyclopedia of Psychology 1–42 (Oxford Univ. Press, 2021).33.Boesch, C., Marchesi, P., Marchesi, N., Fruth, B. & Joulian, F. Is nut cracking in wild chimpanzees a cultural behaviour? J. Hum. Evol. 26, 325–338 (1994).
Google Scholar 
34.Carvalho, S., Matsuzawa, T. & McGrew, W. C. in Tool Use in Animals: Cognition and Ecology (eds Sanz, C. et al.) 225–241 (Cambridge Univ. Press, 2013).35.Morgan, B. & Abwe, E. Chimpanzees use stone hammers in Cameroon. Curr. Biol. 16, 632–633 (2006).
Google Scholar 
36.Sumita, K., Kitahara-Frisch, J. & Norikoshi, K. The aquisition of stone-tool use in captive chimpanzees. Primates 26, 168–181 (1985).
Google Scholar 
37.Marshall-Pescini, S. & Whiten, A. Social learning of nut-cracking behaviour in East African sanctuary-living chimpanzees (Pan troglodytes schweinfurthii). J. Comp. Psychol. 122, 186–194 (2008).PubMed 

Google Scholar 
38.Inoue-Nakamura, N. & Matsuzawa, T. Development of stone tool use by wild chimpanzees (Pan troglodytes). J. Comp. Psychol. 111, 159–173 (1997).CAS 
PubMed 

Google Scholar 
39.van Schaik, C. P., Deaner, R. O. & Merrill, M. The conditions for tool use in primates: implications for the evolution of material culture. J. Hum. Evol. 36, 719–741 (1999).PubMed 

Google Scholar 
40.Humle, T. & Matsuzawa, T. Oil palm use by adjacent communities of chimpanzees at Bossou and Nimba Mountains, West Africa. Int. J. Primatol. 25, 551–581 (2004).
Google Scholar 
41.Koops, K., McGrew, W. C. & Matsuzawa, T. Ecology of culture: do environmental factors influence foraging tool use in wild chimpanzees (Pan troglodytes verus)? Anim. Behav. 85, 175–185 (2013).
Google Scholar 
42.Koops, K., Visalberghi, E. & van Schaik, C. P. The ecology of primate material culture. Biol. Lett. 10, 20140508 (2014).PubMed 
PubMed Central 

Google Scholar 
43.Matsuzawa, T., Humle, T. & Sugiyama, Y. The Chimpanzees of Bossou and Nimba (eds Matsuzawa, T. & Yamagiwa, J.) (Springer, 2011).44.Matsuzawa, T. et al. in Primate Origins of Human Cognition and Behavior (ed. Matsuzawa, T.) 557–574 (Springer, 2001).45.Bandini, E., Motes-Rodrigo, A., Steele, M. P., Rutz, C. & Tennie, C. Examining the mechanisms underlying the acquisition of animal tool behaviour. Biol. Lett. 16, 20200122 (2020).PubMed 
PubMed Central 

Google Scholar 
46.Koops, K., McGrew, W. C., Matsuzawa, T. & Knapp, L. A. Terrestrial nest-building by wild chimpanzees (Pan troglodytes): implications for the tree-to-ground sleep transition in early hominins. Am. J. Phys. Anthropol. 148, 351–361 (2012).PubMed 

Google Scholar 
47.Schuppli, C. et al. The effects of sociability on exploratory tendency and innovation repertoires in wild Sumatran and Bornean orangutans. Sci. Rep. 7, 15464 (2017).PubMed 
PubMed Central 

Google Scholar 
48.Roberts, G. Why individual vigilance declines as group size increases. Anim. Behav. 51, 1077–1086 (1996).
Google Scholar 
49.Visalberghi, E. & Addessi, E. Seeing group members eating a familiar food enhances the acceptance of novel foods in capuchin monkeys. Anim. Behav. 60, 69–76 (2000).CAS 
PubMed 

Google Scholar 
50.Kalan, A. K. et al. Novelty response of wild African apes to camera traps. Curr. Biol. 29, 1211–1217 (2019).CAS 
PubMed 

Google Scholar 
51.Gruber, T., Zuberbühler, K. & Neumann, C. Travel fosters tool use in wild chimpanzees. eLife 5, e16371 (2016).PubMed 
PubMed Central 

Google Scholar 
52.Grund, C., Neumann, C., Zuberbühler, K. & Gruber, T. Necessity creates opportunities for chimpanzee tool use. Behav. Ecol. 30, 1136–1144 (2019).
Google Scholar 
53.Kühl, H. S. et al. Human impact erodes chimpanzee behavioral diversity. Science 363, 1453–1455 (2019).PubMed 

Google Scholar 
54.Wrangham, R. W. Chimpanzees: the culture-zone concept becomes untidy. Curr. Biol. 16, R634–R635 (2006).CAS 
PubMed 

Google Scholar 
55.McGrew, W. C., Ham, R. M., White, L. J. T., Tutin, C. E. G. & Fernandez, M. Why don’t chimpanzees in Gabon crack nuts? Int. J. Primatol. 18, 353–374 (1997).
Google Scholar 
56.Whiten, A. Experimental studies illuminate the cultural transmission of percussive technologies in Homo and Pan. Philos. Trans. R. Soc. B 370, 20140359 (2015).
Google Scholar 
57.Hirata, S., Morimura, N. & Houki, C. How to crack nuts: acquisition process in captive chimpanzees (Pan troglodytes) observing a model. Anim. Cogn. 12, 87–101 (2009).PubMed 

Google Scholar 
58.Hayashi, M., Mizuno, Y. & Matsuzawa, T. How does stone-tool use emerge? Introduction of stones and nuts to naïve chimpanzees in captivity. Primates 46, 91–102 (2005).PubMed 

Google Scholar 
59.Funk, M. Werkzeuggebrauch Beim öffnen von Niessen: Unterschiedliche Bewaltigungen des Problems bei Schimpansen und Orang-Utans (Univ. of Zurich, 1985).60.Visalberghi, E. Acquisition of nut‐cracking behavior by two capuchin monkeys (Cebus apella). Folia Primatol. 49, 168–181 (1987).
Google Scholar 
61.Resende, B. D., Ottoni, E. B. & Fragaszy, D. M. Ontogeny of manipulative behavior and nut-cracking in young tufted capuchin monkeys (Cebus apella): a perception–action perspective. Dev. Sci. 11, 828–840 (2008).PubMed 

Google Scholar 
62.Bandini, E., Grossmann, J., Funk, M., Albiach-Serrano, A. & Tennie, C. Naïve orangutans (Pongo abelii and Pongo pygmaeus) individually acquire nut-cracking using hammer tools. Am. J. Primatol. 83, e23304 (2021).PubMed 

Google Scholar 
63.Damerius, L. A. et al. Orientation toward humans predicts cognitive performance in orang-utans. Sci. Rep. 7, 40052 (2017).CAS 
PubMed 
PubMed Central 

Google Scholar 
64.Reindl, E., Beck, S. R., Apperly, A. & Tennie, C. Young children spontaneously invent wild great apes’ tool-use behaviours. Proc. R. Soc. B 283, 20152402 (2016).CAS 
PubMed 
PubMed Central 

Google Scholar 
65.Neldner, K. et al. A cross-cultural investigation of young children’s spontaneous invention of tool use behaviours. R. Soc. Open Sci. 7, 192240 (2020).PubMed 
PubMed Central 

Google Scholar 
66.Lucas, A. J. et al. The value of teaching increases with tool complexity in cumulative cultural evolution. Proc. R. Soc. B 287, 20201885 (2020).PubMed 
PubMed Central 

Google Scholar 
67.Boesch, C. Teaching among wild chimpanzees. Int. J. Primatol. 41, 530–532 (1991).
Google Scholar 
68.Musgrave, S., Morgan, D., Lonsdorf, E., Mundry, R. & Sanz, C. Tool transfers are a form of teaching among chimpanzees. Sci. Rep. 6, 34783 (2016).CAS 
PubMed 
PubMed Central 

Google Scholar 
69.Koops, K. in The Chimpanzees of Bossou and Nimba (eds Matsuzawa, T. et al.) 277–287 (Springer, 2011).70.van Schaik, C. P. & Pfannes, K. R. in Seasonality in Primates: Studies of Living and Extinct Human and Non-human Primates (eds Brockman, D. K. & van Schaik, C. P.) 23–54 (Cambridge Univ. Press, 2005).71.Matsuzawa, T. in The Chimpanzees of Bossou and Nimba (eds Matsuzawa, T. et al.) 157–164 (Springer, 2011).72.Carvalho, S., Cunha, E., Sousa, C. & Matsuzawa, T. Chaînes opératoires and resource exploitation strategies in chimpanzee nut-cracking (Pan troglodytes). J. Hum. Evol. 55, 148–163 (2008).PubMed 

Google Scholar 
73.Sanz, C., Morgan, D. & Gulick, S. New insights into chimpanzees, tools, and termites from the Congo basin. Am. Nat. 164, 67–581 (2004).
Google Scholar 
74.Hockings, K. J., Anderson, J. R. & Matsuzawa, T. Flexible feeding on cultivated underground storage organs by rainforest-dwelling chimpanzees at Bossou, West Africa. J. Hum. Evol. 58, 227–233 (2010).PubMed 

Google Scholar 
75.Takemoto, H. Seasonal change in terrestriality of chimpanzees in relation to microclimate in the tropical forest. Am. J. Phys. Anthropol. 124, 81–92 (2004).PubMed 

Google Scholar 
76.van Leeuwen, K. L., Matsuzawa, T., Sterck, E. H. M. & Koops, K. How to measure chimpanzee party size? A methodological comparison. Primates 61, 201–212 (2020).PubMed 

Google Scholar 
77.Field, A. P. Discovering Statistics Using IBM SPSS Statistics (SAGE, 2018).78.Humle, T. in The Chimpanzees of Bossou and Nimba (eds Matsuzawa, T. et al.) 61–72 (Springer, 2011).79.Humle, T. & Matsuzawa, T. Behavioural diversity among the wild chimpanzee populations of Bossou and neighbouring areas, Guinea and Côte d’Ivoire, West Africa. Folia Primatol. 72, 57–68 (2001).CAS 

Google Scholar 
80.Sugiyama, Y. & Koman, J. Tool-using and making behavior in wild chimpanzees at Bossou, Guinea. Primates 20, 513–524 (1979).
Google Scholar 
81.Sugiyama, Y. & Koman, J. A preliminary list of chimpanzees’ alimentation at Bossou, Guinea. Primates 28, 133–147 (1987).
Google Scholar 
82.Joulian, F. in Modelling the Early Human Mind (eds Mellars, P. & Gibson, K.) 173–189 (McDonald Institute Monographs, 1996).83.Matsuzawa, T. & Yamakoshi, G. in Reaching into Thought: The Minds of the Great Apes (eds Russon, A. E. et al.) 211–232 (Cambridge Univ. Press, 1996). More

1.Tilman, D., Wedin, D. & Knops, J. Productivity and sustainability influenced by biodiversity in grassland ecosystems. Nature 379, 718–720. https://doi.org/10.1038/379718a0 (1996).ADS 
CAS 
Article 

Google Scholar 
2.Hector, A. et al. Plant diversity and productivity experiments in European grasslands. Science 286, 1123–1127. https://doi.org/10.1126/science.286.5442.1123 (1999).CAS 
Article 
PubMed 

Google Scholar 
3.Kirwan, L. et al. Evenness drives consistent diversity effects in intensive grassland systems across 28 European sites. J. Ecol. 95, 530–539. https://doi.org/10.1111/j.1365-2745.2007.01225.x (2007).Article 

Google Scholar 
4.Sturludóttir, E. et al. Benefits of mixing grasses and legumes for herbage yield and nutritive value in Northern Europe and Canada. Grass Forage Sci. 69, 229–240. https://doi.org/10.1111/gfs.12037 (2014).CAS 
Article 

Google Scholar 
5.Roscher, C. et al. Overyielding in experimental grassland communities – irrespective of species pool or spatial scale. Ecol. Lett. 8, 419–429. https://doi.org/10.1111/j.1461-0248.2005.00736.x (2005).Article 

Google Scholar 
6.Prieto, I. et al. Complementary effects of species and genetic diversity on productivity and stability of sown grasslands. Nat. Plants 1, 1–5. https://doi.org/10.1038/nplants.2015.33 (2015).ADS 
CAS 
Article 

Google Scholar 
7.Fridley, J. D. Resource availability dominates and alters the relationship between species diversity and ecosystem productivity in experimental plant communities. Oecologia 132, 271–277. https://doi.org/10.1007/s00442-002-0965-x (2002).ADS 
Article 
PubMed 

Google Scholar 
8.Sanderson, M. A. et al. Plant species diversity and management of temperate forage and grazing land ecosystems. Crop Sci. 44, 1132–1144. https://doi.org/10.2135/cropsci2004.1132 (2004).Article 

Google Scholar 
9.Hooper, D. U. et al. Effects of biodiversity on ecosystem functioning: a consensus of current knowledge. Ecol. Monogr. 75, 3–35. https://doi.org/10.1890/04-0922 (2005).Article 

Google Scholar 
10.Spehn, E. M. et al. Ecosystem effects of biodiversity manipulations in European grasslands. Ecol. Monogr. 75, 37–63. https://doi.org/10.1890/03-4101 (2005).Article 

Google Scholar 
11.Wright, A. J., Wardle, D. A., Callaway, R. & Gaxiola, A. The overlooked role of facilitation in biodiversity experiments. Trends Ecol. Evol. 32, 383–390. https://doi.org/10.1016/j.tree.2017.02.011 (2017).Article 
PubMed 

Google Scholar 
12.Brophy, C. et al. Major shifts in species’ relative abundance in grassland mixtures alongside positive effects of species diversity in yield: a continental-scale experiment. J. Ecol. 105, 1210–1222. https://doi.org/10.1111/1365-2745.12754 (2017).CAS 
Article 

Google Scholar 
13.Nyfeler, D. et al. Strong mixture effects among four species in fertilized agricultural grassland led to persistent and consistent transgressive overyielding. J. Appl. Ecol. 46, 683–691. https://doi.org/10.1111/j.1365-2664.2009.01653.x (2009).Article 

Google Scholar 
14.Søegaard, K., Gierus, M., Hopkins, A. & Halling, M. Temporary grassland – challenges in the future. Grassl. Sci. Eur. 12, 27–38. https://www.europeangrassland.org/fileadmin/documents/Infos/Printed_Matter/Proceedings/EGF2007_GSE_vol12.pdf (2007).15.Høgh-Jensen, H., Nielsen, B. & Thamsborg, S. M. Productivity and quality, competition and facilitation of chicory in ryegrass/legume-based pastures under various nitrogen supply levels. Eur. J. Agron. 24, 247–256. https://doi.org/10.1016/j.eja.2005.10.007 (2006).CAS 
Article 

Google Scholar 
16.Cong, W. F., Jing, J., Rasmussen, J., Søegaard, K. & Eriksen, J. Forbs enhance productivity of unfertilised grass-clover leys and support low-carbon bioenergy. Sci. Rep. 7, 1–10. https://doi.org/10.1038/s41598-017-01632-4 (2017).CAS 
Article 

Google Scholar 
17.Loreau, M. & Hector, A. Partitioning selection and complementarity in biodiversity experiments. Nature 412, 72–76. https://doi.org/10.1038/35083573 (2001).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
18.Loreau, M., Sapijanskas, J., Isbell, F. & Hector, A. Niche and fitness differences relate the maintenance of diversity to ecosystem function: comment. Ecology 93, 1482–1487. https://doi.org/10.1890/11-0792.1 (2012).Article 
PubMed 

Google Scholar 
19.Montazeaud, G. et al. Crop mixtures: does niche complementarity hold for belowground resources? An experimental test using rice genotypic pairs. Plant Soil 424, 187–202. https://doi.org/10.1007/s11104-017-3496-2 (2018).CAS 
Article 

Google Scholar 
20.Vermeulen, P. J., Van Ruijven, J., Anten, N. P. & Van der Werf, W. An evolutionary game theoretical model shows the limitations of the additive partitioning method for interpreting biodiversity experiments. J. Ecol. 105, 345–353. https://doi.org/10.1111/1365-2745.12706 (2017).Article 

Google Scholar 
21.Hille Ris Lambers, J. H. R., Harpole, W. S., Tilman, D., Knops, J. & Reich, P. B. Mechanisms responsible for the positive diversity–productivity relationship in Minnesota grasslands. Ecol. Lett. 7, 661–668. https://doi.org/10.1111/j.1461-0248.2004.00623.x (2004).22.Marquard, E. et al. Plant species richness and functional composition drive overyielding in a six-year grassland experiment. Ecology 90, 3290–3302. https://doi.org/10.1890/09-0069.1 (2009).Article 
PubMed 

Google Scholar 
23.Lüscher, A., Suter, M., Finn, J., Collins, R. & Gastal, F. Quantification of the effect of legume proportion in the sward on yield advantage and options to keep stable legume proportions (over climatic zones relevant for livestock production). HAL 7, 1–35. https://hal.archives-ouvertes.fr/hal-01611404 (2014).24.Roscher, C., Schumacher, J., Weisser, W. W., Schmid, B. & Schulze, E. D. Detecting the role of individual species for overyielding in experimental grassland communities composed of potentially dominant species. Oecologia 154, 535–549. https://doi.org/10.1007/s00442-007-0846-4 (2007).ADS 
Article 
PubMed 

Google Scholar 
25.Finn, J. A. et al. Ecosystem function enhanced by combining four functional types of plant species in intensively managed grassland mixtures: a 3-year continental-scale field experiment. J. Appl. Ecol. 50, 365–375. https://doi.org/10.1111/1365-2664.12041 (2013).Article 

Google Scholar 
26.Grange, G., Finn, J. A. & Brophy, C. Plant diversity enhanced yield and mitigated drought impacts in intensively managed grassland communities. J. Appl. Ecol. 58, 1864–1875. https://doi.org/10.1111/1365-2664.13894 (2021).CAS 
Article 

Google Scholar 
27.Cardinale, B. J. et al. Impacts of plant diversity on biomass production increase through time because of species complementarity. Proc. Natl. Acad. Sci. U.S.A. 104, 18123–18128. https://doi.org/10.1073/pnas.0709069104 (2007).ADS 
Article 
PubMed 
PubMed Central 

Google Scholar 
28.Marquard, E. et al. Changes in the abundance of grassland species in monocultures versus mixtures and their relation to biodiversity effects. PLoS ONE 8, e75599. https://doi.org/10.1371/journal.pone.0075599 (2013).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
29.Guerrero-Ramírez, N. R. et al. Diversity-dependent temporal divergence of ecosystem functioning in experimental ecosystems. Nat. Ecol. Evol. 1, 1639–1642. https://doi.org/10.1038/s41559-017-0325-1 (2017).Article 
PubMed 
PubMed Central 

Google Scholar 
30.Fargione, J. et al. From selection to complementarity: shifts in the causes of biodiversity-productivity relationships in a long-term biodiversity experiment. Proc. Royal Soc. B 274, 871–876. https://doi.org/10.1098/rspb.2006.0351 (2007).Article 

Google Scholar 
31.Grant, K., Kreyling, J., Heilmeier, H., Beierkuhnlein, C. & Jentsch, A. Extreme weather events and plant-plant interactions: shifts between competition and facilitation among grassland species in the face of drought and heavy rainfall. Ecol. Res. 29, 991–1001. https://doi.org/10.1007/s11284-014-1187-5 (2014).Article 

Google Scholar 
32.Reich, P. B. et al. Impacts of biodiversity loss escalate through time as redundancy fades. Science 336, 589–592. https://doi.org/10.1126/science.1217909 (2012).ADS 
CAS 
Article 
PubMed 

Google Scholar 
33.Roscher, C., Schmid, B., Kolle, O. & Schulze, E. D. Complementarity among four highly productive grassland species depends on resource availability. Oecologia 181, 571–582. https://doi.org/10.1007/s00442-016-3587-4 (2016).ADS 
Article 
PubMed 

Google Scholar 
34.Frankow-Lindberg, B. E., Brophy, C., Collins, R. P. & Connolly, J. Biodiversity effects on yield and unsown species invasion in a temperate forage ecosystem. Ann. Bot. 103, 913–921. https://doi.org/10.1093/aob/mcp008 (2009).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
35.Caradus, J. R. & Woodfield, D. R. World checklist of white clover varieties II. New Zealand J. Agric. Res. 40, 115–206. https://doi.org/10.1080/00288233.1997.9513239 (1997).Article 

Google Scholar 
36.Annicchiarico, P., Barrett, B., Brummer, E. C., Julier, B. & Marshall, A. H. Achievements and challenges in improving temperate perennial forage legumes. Crit. Rev. Plant Sci. 34, 327–380. https://doi.org/10.1080/07352689.2014.898462 (2015).CAS 
Article 

Google Scholar 
37.Abberton, M. T. & Marshall, A. H. Progress in breeding perennial clovers for temperate agriculture. J. Agric. Sci. 143, 117–135. https://doi.org/10.1017/S0021859605005101 (2005).Article 

Google Scholar 
38.Evans, D. R., Williams, T. A. & Mason, S. A. Contribution of white clover varieties to total sward production under typical farm management. Grass Forage Sci. 45, 129–134. https://doi.org/10.1111/j.1365-2494.1990.tb02193.x (1990).Article 

Google Scholar 
39.Hooper, D. U. & Dukes, J. S. Overyielding among plant functional groups in a long-term experiment. Ecol. Lett. 7, 95–105. https://doi.org/10.1046/j.1461-0248.2003.00555.x (2004).Article 

Google Scholar 
40.Ergon, Å. et al. Species interactions in a grassland mixture under low nitrogen fertilization and two cutting frequencies: 1. dry-matter yield and dynamics of species composition. Grass Forage Sci. 71, 667–682. https://doi.org/10.1111/gfs.12250 (2016).CAS 
Article 

Google Scholar 
41.Van Ruijven, J. & Berendse, F. Diversity–productivity relationships: Initial effects, long-term patterns, and underlying mechanisms. Proc. Natl. Acad. Sci. U.S.A. 102, 695–700. https://doi.org/10.1073/pnas.0407524102 (2005).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
42.Barry, K. E. et al. The future of complementarity: disentangling causes from consequences. Trends Ecol. Evol. 34, 167–180. https://doi.org/10.1016/j.tree.2018.10.013 (2019).Article 
PubMed 

Google Scholar 
43.Annicchiarico, P. Breeding white clover for increased ability to compete with associated grasses. J. Agric. Sci. 140, 255–266. https://doi.org/10.1017/S0021859603003198 (2003).Article 

Google Scholar 
44.Zuppinger-Dingley, D. et al. Selection for niche differentiation in plant communities increases biodiversity effects. Nature 515, 108–111. https://doi.org/10.1038/nature13869 (2014).ADS 
CAS 
Article 
PubMed 

Google Scholar 
45.Schöb, C. et al. Intraspecific genetic diversity and composition modify species-level diversity–productivity relationships. New Phytol. 205, 720–730. https://doi.org/10.1111/nph.13043 (2015).Article 
PubMed 

Google Scholar 
46.Barot, S. et al. Designing mixtures of varieties for multifunctional agriculture with the help of ecology A review. Agron. Sustain. Dev. 37, 13. https://doi.org/10.1007/s13593-017-0418-x (2017).Article 

Google Scholar 
47.Van Ruijven, J. & Berendse, F. Long-term persistence of a positive plant diversity–productivity relationship in the absence of legumes. Oikos 118, 101–106. https://doi.org/10.1111/j.1600-0706.2008.17119.x (2009).Article 

Google Scholar 
48.Roscher, C. et al. A functional trait-based approach to understand community assembly and diversity–productivity relationships over 7 years in experimental grasslands. Perspect. Plant Ecol. Evol. Syst. 15, 139–149. https://doi.org/10.1016/j.ppees.2013.02.004 (2013).Article 

Google Scholar 
49.Roscher, C., Thein, S., Schmid, B. & Scherer-Lorenzen, M. Complementary nitrogen use among potentially dominant species in a biodiversity experiment varies between two years. J. Ecol. 96, 477–488. https://doi.org/10.1111/j.1365-2745.2008.01353.x (2008).Article 

Google Scholar 
50.Kayser, M., Müller, J. & Isselstein, J. Grassland renovation has important consequences for C and N cycling and losses. Food Energy Secur. 7, e00146. https://doi.org/10.1002/fes3.146 (2018).Article 

Google Scholar 
51.Bundessortenamt. Beschreibende Sortenliste Futtergräser Esparsette, Klee, Luzerne 2020. https://www.bundessortenamt.de/bsa/media/Files/BSL/bsl_futtergraeser_2020.pdf (2020).52.Rognli, O. A., Pecetti, L., Kovi, M. R. & Annicchiarico, P. Grass and legume breeding matching the future needs of European grassland farming. Grass Forage Sci. 76, 175–185. https://doi.org/10.1111/gfs.12535 (2021).Article 

Google Scholar 
53.Annicchiarico, P. et al. Do we need specific breeding for legume-based mixtures?. Adv. Agron. 157, 141–215. https://doi.org/10.1016/bs.agron.2019.04.001 (2019).Article 

Google Scholar 
54.Sampoux, J. P., Giraud, H. & Litrico, I. Which recurrent selection scheme to improve mixtures of crop species? Theoretical expectations. G3-Genes Genom. Genet. 10, 89–107. https://doi.org/10.1534/g3.120.401092 (2020).55.Hofer, D. et al. Yield of temperate forage grassland species is either largely resistant or resilient to experimental summer drought. J. Appl. Ecol. 53, 1023–1034. https://doi.org/10.1111/1365-2664.12694 (2016).Article 

Google Scholar 
56.Pinheiro, J. et al. Nlme: Linear and nonlinear mixed effects models. R package version 3.1–137. https://CRAN.R-project.org/package=nlme (2018).57.Bartón, K. MuMIn: Multi-model inference. R package version 1.43.6. https://CRAN.R-project.org/package=MuMIn (2019). More

1.Harley, C. D. G. et al. The impacts of climate change in coastal marine systems. Ecol. Lett. 9, 228–241 (2006).ADS 
PubMed 

Google Scholar 
2.Poloczanska, E. S. et al. Global imprint of climate change on marine life. Nat. Clim. Change 3, 919–925 (2013).ADS 

Google Scholar 
3.Hoegh-Guldberg, O. et al. Impacts of 1.5 °C global warming on natural and human systems. In: […]. in Global Warming of 1.5 °C. An IPCC Special Report on the Impacts of Global Warming of 1.5 °C Above Preindustrial Levels and Related Global Greenhouse Gas Emission Pathways, in the Context of Strengthening the Global Response to the Threat of Climate Change (eds. Masson-Delmotte, V. et al.) 175–311 (2018).4.Gunderson, A. R., Armstrong, E. J. & Stillman, J. H. Multiple stressors in a changing world: The need for an improved perspective on physiological responses to the dynamic marine environment. Ann. Rev. Mar. Sci. 8, 357–378 (2016).PubMed 

Google Scholar 
5.Ban, S. S., Graham, N. A. J. & Connolly, S. R. Evidence for multiple stressor interactions and effects on coral reefs. Glob. Change Biol. 20, 681–697 (2014).ADS 

Google Scholar 
6.Marcogliese, D. J. The impact of climate change on the parasites and infectious diseases of aquatic animals. OIE Rev. Sci. Tech. 27, 467–484 (2008).CAS 

Google Scholar 
7.Urban, M. C., Tewksbury, J. J. & Sheldon, K. S. On a collision course: Competition and dispersal differences create no-analogue communities and cause extinctions during climate change. Proc. R. Soc. B Biol. Sci. 279, 2072–2080 (2012).
Google Scholar 
8.Mouritsen, K. N. & Poulin, R. Parasitism, climate oscillations and the structure of natural communities. Oikos 97, 462–468 (2002).
Google Scholar 
9.Poulin, R. & Mouritsen, K. N. Climate change, parasitism and the structure of intertidal ecosystems. J. Helminthol. 80, 183–191 (2006).CAS 
PubMed 

Google Scholar 
10.Mouritsen, K. N., Sørensen, M. M., Poulin, R. & Fredensborg, B. L. Coastal ecosystems on a tipping point: Global warming and parasitism combine to alter community structure and function. Glob. Change Biol. 24, 4340–4356 (2018).ADS 

Google Scholar 
11.James, C. C. et al. Marine host–pathogen dynamics: Influences of global climate change. Oceanography 31, 182–193 (2018).
Google Scholar 
12.Friesen, O. C., Poulin, R. & Lagrue, C. Temperature and multiple parasites combine to alter host community structure. Oikos 130(9), 1500–1511 (2021).

Google Scholar 
13.Poulin, R. Parasite biodiversity revisited: Frontiers and constraints. Int. J. Parasitol. 44, 581–589 (2014).PubMed 

Google Scholar 
14.Kuris, A. M. et al. Ecosystem energetic implications of parasite and free-living biomass in three estuaries. Nature 454, 515–518 (2008).ADS 
CAS 
PubMed 

Google Scholar 
15.Galaktionov, K. V. & Dobrovolskij, A. A. The Biology and Evolution of Trematodes: An Essay on the Biology, Morphology, Life Cycles, Transmissions, and Evolution of Digenetic Trematodes. The American Journal of Semiotics vol. 4 (Springer-Science+Business Media Dordrecht, 2003).16.Thieltges, D. W., Jensen, K. T. & Poulin, R. The role of biotic factors in the transmission of free-living endohelminth stages. Parasitology 135, 407–426 (2008).CAS 
PubMed 

Google Scholar 
17.Pietrock, M. & Marcogliese, D. J. Free-living endohelminth stages: At the mercy of environmental conditions. Trends Parasitol. 19, 293–299 (2003).PubMed 

Google Scholar 
18.Morley, N. J. Thermodynamics of cercarial survival and metabolism in a changing climate. Parasitology 138, 1442–1452 (2011).CAS 
PubMed 

Google Scholar 
19.Poulin, R. Global warming and temperature-mediated increases in cercarial emergence in trematode parasites. Parasitology 132, 143–151 (2006).CAS 
PubMed 

Google Scholar 
20.Thieltges, D. W. & Rick, J. Effect of temperature on emergence, survival and infectivity of cercariae of the marine trematode Renicola roscovita (Digenea: Renicolidae). Dis. Aquat. Organ. 73, 63–68 (2006).PubMed 

Google Scholar 
21.Selbach, C. & Poulin, R. Some like it hotter: Trematode transmission under changing temperature conditions. Oecologia 194, 745–755 (2020).ADS 
PubMed 

Google Scholar 
22.Morley, N. J. Inbred laboratory cultures and natural trematode transmission under climate change. Trends Parasitol. 27, 286–287 (2011).PubMed 

Google Scholar 
23.Paull, S. H. & Johnson, P. T. J. Experimental warming drives a seasonal shift in the timing of host–parasite dynamics with consequences for disease risk. Ecol. Lett. 17, 445–453 (2014).PubMed 

Google Scholar 
24.Paull, S. H., Lafonte, B. E. & Johnson, P. T. J. Temperature-driven shifts in a host–parasite interaction drive nonlinear changes in disease risk. Glob. Change Biol. 18, 3558–3567 (2012).ADS 

Google Scholar 
25.Studer, A., Poulin, R. & Tompkins, D. M. Local effects of a global problem: Modelling the risk of parasite-induced mortality in an intertidal trematode-amphipod system. Oecologia 172, 1213–1222 (2013).ADS 
CAS 
PubMed 

Google Scholar 
26.Studer, A., Thieltges, D. W. & Poulin, R. Parasites and global warming: Net effects of temperature on an intertidal host–parasite system. Mar. Ecol. Prog. Ser. 415, 11–22 (2010).ADS 

Google Scholar 
27.Mouritsen, K. N., Tompkins, D. M. & Poulin, R. Climate warming may cause a parasite-induced collapse in coastal amphipod populations. Oecologia 146, 476–483 (2005).ADS 
PubMed 

Google Scholar 
28.Selbach, C., Barsøe, M., Vogensen, T. K., Samsing, A. B. & Mouritsen, K. N. Temperature–parasite interaction: Do trematode infections protect against heat stress?. Int. J. Parasitol. 50, 1189–1194 (2020).PubMed 

Google Scholar 
29.Werding, B. Morphologie, Entwicklung und Ökologie digener Trematoden-Larven der Strandschnecke Littorina littorea. Mar. Biol. 3, 306–333 (1969).
Google Scholar 
30.Somero, G. N. The physiology of climate change: How potentials for acclimatization and genetic adaptation will determine ‘winners’ and ‘losers’. J. Exp. Biol. 213, 912–920 (2010).CAS 
PubMed 

Google Scholar 
31.Fredensborg, B. L., Mouritsen, K. N. & Poulin, R. Impact of trematodes on host survival and population density in the intertidal gastropod Zeacumantus subcarcinatus. Mar. Ecol. Prog. Ser. 290, 109–117 (2005).ADS 

Google Scholar 
32.Morón Lugo, S. C. et al. Warming and temperature variability determine the performance of two invertebrate predators. Sci. Rep. 10, 1–14 (2020).ADS 

Google Scholar 
33.Wolf, F. et al. High resolution water temperature data between January 1997 and December 2018 at the GEOMAR pier surface. Bremen PANGAEA. https://doi.org/10.1594/PANGAEA.919186 (2020).34.Franz, M., Lieberum, C., Bock, G. & Karez, R. Environmental parameters of shallow water habitats in the SW Baltic Sea. Earth Syst. Sci. Data 11, 947–957 (2019).ADS 

Google Scholar 
35.Lajeunesse, M. J. Bias and correction for the log response ratio in ecological meta-analysis. Ecology 96, 2056–2063 (2015).PubMed 

Google Scholar 
36.Gräwe, U., Friedland, R. & Burchard, H. The future of the western Baltic Sea: Two possible scenarios. Ocean Dyn. 63, 901–921 (2013).ADS 

Google Scholar 
37.Pansch, C. et al. Heat waves and their significance for a temperate benthic community: A near-natural experimental approach. Glob. Change Biol. 24, 4357–4367 (2018).ADS 

Google Scholar 
38.Sokolova, I. M., Frederich, M., Bagwe, R., Lannig, G. & Sukhotin, A. A. Energy homeostasis as an integrative tool for assessing limits of environmental stress tolerance in aquatic invertebrates. Mar. Environ. Res. 79, 1–15 (2012).CAS 
PubMed 

Google Scholar 
39.Clarke, A. P., Mill, P. J. & Grahame, J. The nature of heat coma in Littorina littorea (Mollusca: Gastropoda). Mar. Biol. 137, 447–451 (2000).
Google Scholar 
40.McDaniel, S. J. Littorina littorea: Lowered heat tolerance due to Cryptocotyle lingua. Exp. Parasitol. 25, 13–15 (1969).CAS 
PubMed 

Google Scholar 
41.Ataev, G. Temperature influence on the development and biology of rediae and cercariae of Philophthalmus rhionica (Trematoda). Parazitologiâ 25, 349–359 (1991).
Google Scholar 
42.Paull, S. H. & Johnson, P. T. J. High temperature enhances host pathology in a snail-trematode system: Possible consequences of climate change for the emergence of disease. Freshw. Biol. 56, 767–778 (2011).
Google Scholar 
43.Paull, S. H., Raffel, T. R., Lafonte, B. E. & Johnson, P. T. J. How temperature shifts affect parasite production: Testing the roles of thermal stress and acclimation. Funct. Ecol. 29, 941–950 (2015).
Google Scholar 
44.Kuris, A. M. Effect of exposure to Echinostoma liei miracidia on growth and survival of young Biomphalaria glabrata snails. Int. J. Parasitol. 10, 303–308 (1980).CAS 
PubMed 

Google Scholar 
45.Mouritsen, K. N. & Haun, S. C. B. Community regulation by herbivore parasitism and density: Trait-mediated indirect interactions in the intertidal. J. Exp. Mar. Biol. Ecol. 367, 236–246 (2008).
Google Scholar 
46.Bommarito, C. et al. Effects of first intermediate host density, host size and salinity on trematode infections in mussels of the south-western Baltic Sea. Parasitology 148, 486–494 (2021).CAS 
PubMed 

Google Scholar 
47.McCarthy, A. M. The influence of temperature on the survival and infectivity of the cercariae of Echinoparyphium recurvatum (Digenea: Echinostomatidae). Parasitology 118, 383–388 (1999).PubMed 

Google Scholar 
48.Morley, N. J. & Lewis, J. W. Thermodynamics of trematode infectivity. Parasitology 142, 585–597 (2015).CAS 
PubMed 

Google Scholar 
49.Mouritsen, K. N. & Jensen, K. T. Parasite transmission between soft-bottom invertebrates: Temperature mediated infection rates and mortality in Corophium volutator. Mar. Ecol. Prog. Ser. 151, 123–134 (1997).ADS 

Google Scholar 
50.de Montaudouin, X., Wegeberg, A. M., Jensen, K. T. & Sauriau, P. G. Infection characteristics of Himasthla elongata cercariae in cockles as a function of water current. Dis. Aquat. Organ. 34, 63–70 (1998).
Google Scholar 
51.Vajedsamiei, J. et al. Simultaneous recording of filtration and respiration in marine organisms in response to short-term environmental variability. Limnol. Oceanogr. Methods https://doi.org/10.1002/lom3.10414 (2021).Article 

Google Scholar 
52.Koehler, A. V., Brown, B., Poulin, R., Thieltges, D. W. & Fredensborg, B. L. Disentangling phylogenetic constraints from selective forces in the evolution of trematode transmission stages. Evol. Ecol. 26, 1497–1512 (2012).
Google Scholar 
53.Stunkard, H. W. The morphology and life history of the digenetic trematode, Himasthla littorinae sp. n. (Echinostomatidae). J. Parasitol. 52, 367–372 (2014).
Google Scholar 
54.Selbach, C. & Poulin, R. Parasites in space and time: A novel method to assess and illustrate host-searching behaviour of trematode cercariae. Parasitology 145, 1469–1474 (2018).PubMed 

Google Scholar 
55.Gorbushin, A. M. & Levakin, I. A. Encystment in vitro of the cercariae Himasthla elongata (Trematoda: Echinostomatidae). J. Evol. Biochem. Physiol. 41, 428–436 (2005).
Google Scholar 
56.Gorbushin, A. M. & Shaposhnikova, T. G. In vitro culture of the avian echinostome Himasthla elongata: From redia to marita. Exp. Parasitol. 101, 234–239 (2002).CAS 
PubMed 

Google Scholar 
57.Levakin, I. A., Losev, E. A., Nikolaev, K. E. & Galaktionov, K. V. In vitro encystment of Himasthla elongata cercariae (Digenea, Echinostomatidae) in the haemolymph of blue mussels Mytilus edulis as a tool for assessing cercarial infectivity and molluscan susceptibility. J. Helminthol. 87, 180–188 (2013).CAS 
PubMed 

Google Scholar 
58.Choisy, M., Brown, S. P., Lafferty, K. D. & Thomas, F. Evolution of trophic transmission in parasites: Why add intermediate hosts?. Am. Nat. 162, 172–181 (2003).PubMed 

Google Scholar 
59.Pechenik, J. & Fried, B. Effect of temperature on survival and infectivity of Echinostoma trivolvis cercariae: A test of the energy limitation hypothesis. Parasitology 111, 373–378 (1995).
Google Scholar 
60.Fried, B. & Ponder, E. L. Effects of temperature on survival, infectivity and in vitro encystment of the cercariae of Echinostoma caproni. J. Helminthol. 77, 235–238 (2003).CAS 
PubMed 

Google Scholar 
61.Bommarito, C. et al. Freshening rather than warming drives trematode transmission from periwinkles to mussels. Mar. Biol. 167, 1–12 (2020).
Google Scholar 
62.Morley, N. J. & Lewis, J. W. Thermodynamics of cercarial development and emergence in trematodes. Parasitology 140, 121–1214 (2013).
Google Scholar 
63.Büttger, H. et al. Community dynamics of intertidal soft-bottom mussel beds over two decades. Helgol. Mar. Res. 62, 23–36 (2008).ADS 

Google Scholar 
64.Jaatinen, K., Westerbom, M., Norkko, A., Mustonen, O. & Koons, D. N. Detrimental impacts of climate change may be exacerbated by density-dependent population regulation in blue mussels. J. Anim. Ecol. 90, 562–573 (2021).PubMed 

Google Scholar 
65.Studer, A. & Poulin, R. Analysis of trait mean and variability versus temperature in trematode cercariae: Is there scope for adaptation to global warming?. Int. J. Parasitol. 44, 403–413 (2014).CAS 
PubMed 

Google Scholar 
66.Berkhout, B. W., Lloyd, M. M., Poulin, R. & Studer, A. Variation among genotypes in responses to increasing temperature in a marine parasite: Evolutionary potential in the face of global warming?. Int. J. Parasitol. 44, 1019–1027 (2014).PubMed 

Google Scholar 
67.Vanoverschelde, R. Studies on the life-cycle of Himasthla militaris (Trematoda: Echinostomatidae): Influence of salinity and temperature on egg development and miracidial emergence. Parasitology 82, 459–465 (1981).
Google Scholar 
68.Vanoverschelde, R. Studies on the life-cycle of Himasthla militaris (Trematoda: Echinostomatidae): Influence of temperature and salinity on the life-span of the miracidium and the infection of the first intermediate host, Hydrobia ventrosa. Parasitology 84, 131–135 (1982).
Google Scholar 
69.de Montaudouin, X. et al. Digenean trematode species in the cockle Cerastoderma edule: Identification key and distribution along the North-Eastern Atlantic Shoreline. J. Mar. Biol. Assoc. U.K. 89, 543–556 (2009).
Google Scholar 
70.Richard, A., de Montaudouin, X., Rubiello, A. & Maire, O. Cockle as second intermediate host of trematode parasites: Consequences for sediment bioturbation and nutrient fluxes across the benthic interface. J. Mar. Sci. Eng. 9, 749 (2021).
Google Scholar 
71.Magalhães, L., Freitas, R. & de Montaudouin, X. How costly are metacercarial infections in a bivalve host? Effects of two trematode species on biochemical performance of cockles. J. Invertebr. Pathol. 177, 107479 (2020).PubMed 

Google Scholar 
72.Magalhães, L., de Montaudouin, X., Figueira, E. & Freitas, R. Trematode infection modulates cockles biochemical response to climate change. Sci. Total Environ. 637–638, 30–40 (2018).ADS 
PubMed 

Google Scholar 
73.Magalhães, L. et al. Seasonal variation of transcriptomic and biochemical parameters of cockles (Cerastoderma edule) related to their infection by trematode parasites. J. Invertebr. Pathol. 148, 73–80 (2017).PubMed 

Google Scholar 
74.Bakhmet, I., Nikolaev, K. & Levakin, I. Effect of infection with Metacercariae of Himasthla elongata (Trematoda: Echinostomatidae) on cardiac activity and growth rate in blue mussels (Mytilus edulis) in situ. J. Sea Res. 123, 51–54 (2017).ADS 

Google Scholar 
75.Stier, T., Drent, J. & Thieltges, D. W. Trematode infections reduce clearance rates and condition in blue mussels Mytilus edulis. Mar. Ecol. Prog. Ser. 529, 137–144 (2015).ADS 

Google Scholar 
76.de Montaudouin, X., Bazairi, H. & Culloty, S. Effect of trematode parasites on cockle Cerastoderma edule growth and condition index: A transplant experiment. Mar. Ecol. Prog. Ser. 471, 111–121 (2012).ADS 

Google Scholar 
77.Seuront, L., Nicastro, K. R., Zardi, G. I. & Goberville, E. Decreased thermal tolerance under recurrent heat stress conditions explains summer mass mortality of the blue mussel Mytilus edulis. Sci. Rep. 9, 1–14 (2019).ADS 

Google Scholar 
78.Österblom, H. et al. Human-induced trophic cascades and ecological regime shifts in the baltic sea. Ecosystems 10, 877–889 (2007).
Google Scholar 
79.Zander, C. D. & Reimer, L. W. Parasitism at the ecosystem level in the Baltic Sea. Parasitology 124, 119–135 (2002).
Google Scholar 
80.Johnson, P. T. J. et al. Aquatic eutrophication promotes pathogenic infection in amphibians. Proc. Natl. Acad. Sci. U. S. A. 104, 15781–15786 (2007).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
81.Budria, A. & Candolin, U. How does human-induced environmental change influence host–parasite interactions?. Parasitology 141, 462–474 (2014).PubMed 

Google Scholar 
82.Aalto, S. L., Decaestecker, E. & Pulkkinen, K. A three-way perspective of stoichiometric changes on host–parasite interactions. Trends Parasitol. 31, 333–340 (2015).PubMed 

Google Scholar 
83.Vajedsamiei, J., Melzner, F., Raatz, M., Moron, S. & Pansch, C. Cyclic thermal fluctuations can be burden or relief for an ectotherm depending on fluctuations’ average and amplitude. Funct. Ecol. 35, 2483–2496 (2021).
Google Scholar 
84.Moisez, E., Spilmont, N. & Seuront, L. Microhabitats choice in intertidal gastropods is species-, temperature- and habitat-specific. J. Therm. Biol. 94, 102785 (2020).PubMed 

Google Scholar 
85.Bates, A. E., Leiterer, F., Wiedeback, M. L. & Poulin, R. Parasitized snails take the heat: A case of host manipulation?. Oecologia 167, 613–621 (2011).ADS 
CAS 
PubMed 

Google Scholar 
86.Shinagawa, K., Urabe, M. & Nagoshi, M. Relationships between trematode infection and habitat depth in a freshwater snail, Semisulcospira libertina (Gould). Hydrobiologia 397, 171–178 (1999).
Google Scholar 
87.Friesen, O. C., Poulin, R. & Lagrue, C. Parasite-mediated microhabitat segregation between congeneric hosts. Biol. Lett. 14, 20170671 (2018).PubMed 
PubMed Central 

Google Scholar 
88.Welsh, J. E., Van Der Meer, J., Brussaard, C. P. D. & Thieltges, D. W. Inventory of organisms interfering with transmission of a marine trematode. J. Mar. Biol. Assoc. U.K. 94, 697–702 (2014).
Google Scholar 
89.Soldánová, M., Selbach, C. & Sures, B. The early worm catches the bird? Productivity and patterns of Trichobilharzia szidati cercarial emission from Lymnaea stagnalis. PLoS One 11, 1–21 (2016).
Google Scholar 
90.Solovyeva, A. et al. Reduced infectivity in Himasthla elongata (Trematoda, Himasthlidae) cercariae with deviant photoreaction. J. Helminthol. 94, 1–5 (2020).
Google Scholar 
91.de Montaudouin, X., Blanchet, H., Desclaux-Marchand, C., Lavesque, N. & Bachelet, G. Cockle infection by Himasthla quissetensis—I. From cercariae emergence to metacercariae infection. J. Sea Res. 113, 99–107 (2016).
Google Scholar 
92.Zuur, A. F., Ieno, E. N., Walker, N. J., Saveliev, A. A. & Smith, G. M. Mixed Effects Models and Extensions in Ecology with R. Statistics for Biology and Health vol. 36 (Springer, 2009).93.Brooks, M. E. et al. glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. R J. 9, 378–400 (2017).
Google Scholar 
94.Hartig, F. DHARMa: Residual Diagnostics for Hierarchical (Multi-Level/Mixed) Regression Models. R package version 0.2.0. (2018).
https://CRAN.R-project.org/package=DHARMa 1–36 https://cran.r-project.org/web/packages/DHARMa/DHARMa.pdf. Accessed 26 Feb 2021.95.Wood, S. N. Generalized Additive Models: An Introduction with R 2nd edn. (CRC, 2017).MATH 

Google Scholar 
96.Wood, S. N., Pya, N. & Säfken, B. Smoothing parameter and model selection for general smooth models. J. Am. Stat. Assoc. 111, 1548–1563 (2016).MathSciNet 
CAS 

Google Scholar 
97.Nakagawa, S. & Schielzeth, H. A general and simple method for obtaining R2 from generalized linear mixed-effects models. Methods Ecol. Evol. 4, 133–142 (2013).
Google Scholar 
98.Nakagawa, S., Johnson, P. C. D. & Schielzeth, H. The coefficient of determination R2 and intra-class correlation coefficient from generalized linear mixed-effects models revisited and expanded. J. R. Soc. Interface 14, 1–11 (2017).
Google Scholar 
99.Lüdecke, D., Makowski, D., Waggoner, P. & Patil, I. Assessment of Regression Models Performance. CRAN. CRAN https://easystats.github.io/performance/ (2020) https://doi.org/10.1098/rsif.2017.0213. Accessed 1 Sept 2021.100.Fox, J. & Weisberg, S. An R Companion to Applied Regression. Robust Regression in R (Sage, 2019).
Google Scholar 
101.Hedges, L. V., Gurevitch, J. & Curtis, P. S. The meta-analysis of response ratios in experimental ecology. Ecology 80, 1150–1156 (1999).
Google Scholar  More

1.Science Advice for Policy by European Academies. A scientific perspective on microplastics in nature and society (SAPEA, 2019). Expert group report summarizing the state of the science regarding microplastics in nature and society.2.Koelmans, A. A. et al. Risks of plastic debris: Unravelling fact, opinion, perception and belief. Environ. Sci. Technol. 51, 11513–11519 (2017).CAS 

Google Scholar 
3.Henderson, L. & Green, C. Making sense of microplastics? Public understandings of plastic pollution. Mar. Pollut. Bull. 152, 110908 (2020).CAS 

Google Scholar 
4.Group of Experts on the Scientific Aspects of Marine Environmental Protection (GESAMP). Sources, fate and effects of microplastics in the marine environment. Part two of a global assessment (eds Kershaw, P. J. & Rochman, C. M.) (IMO/FAO/UNESCO-IOC/UNIDO/WMO/IAEA/UN/UNEP/UNDP, 2016).5.Arthur, C., Baker, J. & Bamford, H. (eds) NOAA technical memorandum NOS-OR&R-30. In Proc. Int. Res. Worksh. Occurrence, Effects and Fate of Microplastic Marine Debris (NOAA, 2009).6.European Chemicals Agency. Annex XV restriction report proposal for a restriction: intentionally added microplastics. Version 1.2. Proposal 1.2. ECA https://echa.europa.eu/documents/10162/05bd96e3-b969-0a7c-c6d0-441182893720 (2019).7.Coffin, S. Proposed definition of ‘microplastics in drinking water’. California Water Boards https://www.waterboards.ca.gov/drinking_water/certlic/drinkingwater/docs/stffrprt_jun3.pdf (2020).8.Andrady, A. L. Microplastics in the marine environment. Mar. Pollut. Bull. 62, 1596–1605 (2011).CAS 

Google Scholar 
9.Kooi, M. & Koelmans, A. A. Simplifying microplastic via continuous probability distributions for size, shape and density. Environ. Sci. Technol. Lett. 6, 551–557 (2019). This paper introduces the concept of describing microplastic characteristics through continuous PDFs, allowing us to capture the diversity of microplastics as a single contaminant in transport, exposure and risk assessment, rather than across many separate categories.CAS 

Google Scholar 
10.Rochman, C. M. et al. Rethinking microplastics as a diverse contaminant suite. Environ. Toxicol. Chem. 38, 703–711 (2019).CAS 

Google Scholar 
11.Kooi, M., Besseling, E., Kroeze, C., van Wezel, A. P. & Koelmans, A. A. Modelling the fate and transport of plastic debris in fresh waters. Review and guidance. In Freshwater Microplastics. The Handbook of Environmental Chemistry Vol. 58 (eds Wagner M. & Lambert S.) 125–152 (Springer, 2017).12.Hardesty, B. D. et al. Using numerical model simulations to improve the understanding of micro-plastic distribution and pathways in the marine environment. Front. Mar. Sci. 4, 1–30 (2017).
Google Scholar 
13.Redondo-Hasselerharm, P. E., Falahudin, D., Peeters, E. T. H. M. & Koelmans, A. A. Microplastic effect thresholds for freshwater benthic macroinvertebrates. Environ. Sci. Technol. 52, 2278–2286 (2018).CAS 

Google Scholar 
14.Adam, V., Yang, T. & Nowack, B. Toward an ecotoxicological risk assessment of microplastics: comparison of available hazard and exposure data in freshwaters. Environ. Toxicol. Chem. 38, 436–447 (2019). This paper introduces probabilistic SSDs for microplastic particles.CAS 

Google Scholar 
15.Koelmans, A. A., Diepens N. J. & Mohamed Nor, N. H. Weight of evidence for the microplastic vector effect in the context of chemical risk assessment. In Microplastic in the Environment: Pattern and Process (ed. Bank, M. S.) (Springer, 2021).16.Besseling, E., Redondo-Hasselerharm, P. E., Foekema, E. M. & Koelmans, A. A. Quantifying ecological risks of aquatic micro- and nanoplastic. Crit. Rev. Environ. Sci. Technol. 49, 32–80 (2019).
Google Scholar 
17.Burns, E. E. & Boxall, A. B. A. Microplastics in the aquatic environment: evidence for or against adverse impacts and major knowledge gaps. Environ. Toxicol. Chem. 37, 2776–2796 (2018).CAS 

Google Scholar 
18.Wright, S. L. & Kelly, F. J. Plastic and human health: a micro issue? Environ. Sci. Technol. 51, 6634–6647 (2017). This is a thorough review and outlook on the implications of plastic for human health.CAS 

Google Scholar 
19.Mohamed Nor, N. H., Kooi, M., Diepens, N. J. & Koelmans, A. A. Lifetime accumulation of nano- and microplastic in children and adults. Environ. Sci. Technol. 55, 5084–5096 (2021). This paper is the first probabilistic and aligned microplastic exposure assessment for humans, using PDFs.CAS 

Google Scholar 
20.Noventa, S. et al. Paradigms to assess the human health risks of nano- and microplastics. Micropl. Nanopl. 1, 9 (2021).
Google Scholar 
21.Ramsperger, A. F. R. M. et al. Environmental exposure enhances the internalization of microplastic particles into cells. Sci. Adv. 6, eabd1211 (2020).CAS 

Google Scholar 
22.Rejman, J., Oberle, V., Zuhorn, I. S. & Hoekstra, D. Size-dependent internalization of particles via the pathways of clathrin- and caveolae-mediated endocytosis. Biochem. J. 377, 159–169 (2004).CAS 

Google Scholar 
23.Ragusa, A. et al. Plasticenta: first evidence of microplastics in human placenta. Environ. Int. 146, 106274 (2021).CAS 

Google Scholar 
24.Schwabl, P. et al. Detection of various microplastics in human stool: a prospective case series. Ann. Intern. Med. 171, 453–457 (2019).
Google Scholar 
25.Connors, K. A., Dyer, S. D. & Belanger, S. E. Advancing the quality of environmental microplastic research. Environ. Toxicol. Chem. 36, 1697–1703 (2017). This paper highlights the need for better quality in microplastic research.CAS 

Google Scholar 
26.Wesch, C., Bredimus, K., Paulus, M. & Klein, R. Towards the suitable monitoring of ingestion of microplastics by marine biota: a review. Environ. Pollut. 218, 1200–1208 (2016).CAS 

Google Scholar 
27.O’Connor, J. et al. Microplastics in freshwater biota: a critical review of isolation, characterization and assessment methods. Glob. Challeng. https://doi.org/10.1002/gch2.201800118 (2019).28.de Ruijter, V. N., Redondo-Hasselerharm, P. E., Gouin, T. & Koelmans, A. A. Quality criteria for microplastic effect studies in the context of risk assessment: a critical review. Environ. Sci. Technol. 54, 11692–11705 (2020).
Google Scholar 
29.Hartmann, N. B. et al. Are we speaking the same language? Recommendations for a definition and categorization framework for plastic debris. Environ. Sci. Technol. 53, 1039–1047 (2019).CAS 

Google Scholar 
30.Kögel, T., Bjorøy, Ø., Toto, B., Bienfait, A. M. & Sanden, M. Micro- and nanoplastic toxicity on aquatic life: determining factors. Sci. Total. Environ. 709, 136050 (2020).
Google Scholar 
31.Bond, T., Ferrandiz-Mas, V., Felipe-Sotelo, M. & van Sebille, E. The occurrence and degradation of aquatic plastic litter based on polymer physicochemical properties: a review. Crit. Rev. Environ. Sci. Technol. 48, 685 (2018).
Google Scholar 
32.Riediker, M. et al. Particle toxicology and health — where are we? Part. Fibre Toxicol. 16, 1–33 (2019).
Google Scholar 
33.Kooi, M. et al. Characterizing the multidimensionality of microplastics across environmental compartments. Water Res. 202, 117429 (2021).CAS 

Google Scholar 
34.Wiesinger, H., Wang, Z. & Hellweg, S. Deep dive into plastic monomers, additives, and processing aids. Environ. Sci. Technol. 55, 9339–9351 (2021).CAS 

Google Scholar 
35.Gouin, T. Addressing the importance of microplastic particles as vectors for long-range transport of chemical contaminants: perspective in relation to prioritizing research and regulatory actions. Micropl. Nanopl. 1, 14 (2021).
Google Scholar 
36.Hermabessiere, L. et al. Occurrence and effects of plastic additives on marine environments and organisms: a review. Chemosphere 182, 781–793 (2017).CAS 

Google Scholar 
37.Gouin, T., Roche, N., Lohmann, R. & Hodges, G. A thermodynamic approach for assessing the environmental exposure of chemicals absorbed to microplastic. Environ. Sci. Technol. 45, 1466–1472 (2011).CAS 

Google Scholar 
38.Lohmann, R. Microplastics are not important for the cycling and bioaccumulation of organic pollutants in the oceans — but should microplastics be considered POPs themselves? Int. Environ. Assess. Manag. 13, 460–465 (2017).CAS 

Google Scholar 
39.Takada, H. & Karapanagioti, H. K. (eds) Hazardous Chemicals Associated with Plastics in the Marine Environment (Springer International Publishing, 2016).40.Hong, S. H., Shim, W. J. & Hong, K. Methods of analysing chemicals associated with microplastics: a review. Anal. Methods 9, 1361–1368 (2017).
Google Scholar 
41.Koelmans, A. A., Bakir, A., Burton, G. A. & Janssen, C. R. Microplastic as a vector for chemicals in the aquatic environment. critical review and model-supported re-interpretation of empirical studies. Environ. Sci. Technol. 50, 3315–3326 (2016).CAS 

Google Scholar 
42.Jahnke, A. et al. Reducing uncertainty and confronting ignorance about the possible impacts of weathering plastic in the marine environment. Environ. Sci. Technol. Lett. 4, 85–90 (2017).CAS 

Google Scholar 
43.Boucher, J. and Friot D. Primary Microplastics in the Oceans: A Global Evaluation of Sources 43 (IUCN, 2017).44.Koelmans, A. A., Kooi, M., Lavender-Law, K. & Van Sebille, E. All is not lost: deriving a top-down mass budget of plastic at sea. Environ. Res. Lett. 12, 114028 (2017).
Google Scholar 
45.Kawecki, D. & Nowack, D. Polymer-specific modeling of the environmental emissions of seven commodity plastics as macro- and microplastics. Environ. Sci. Technol. 53, 9664–9676 (2019).CAS 

Google Scholar 
46.Kooi, M., Van Nes, E. H., Scheffer, M. & Koelmans, A. A. Ups and downs in the ocean: effects of biofouling on vertical transport of microplastics. Environ. Sci. Technol. 51, 7963–7971 (2017).CAS 

Google Scholar 
47.Mateos-Cárdenas, A., O’Halloran, J., van Pelt, F. N. A. M. & Jansen, M. A. K. Rapid fragmentation of microplastics by the freshwater amphipod Gammarus duebeni (Lillj.). Sci. Rep. 10, 12799 (2020).
Google Scholar 
48.Julienne, F., Delorme, N. & Lagarde, F. From macroplastics to microplastics: role of water in the fragmentation of polyethylene. Chemosphere 236, 124409 (2019).CAS 

Google Scholar 
49.Koelmans, A. A., Redondo-Hasselerharm, P. E., Mohamed Nor, N. H. & Kooi, M. Solving the non-alignment of methods and approaches used in microplastic research in order to consistently characterize risk. Environ. Sci. Technol. 54, 12307–12315 (2020).CAS 

Google Scholar 
50.Cózar, A. et al. Plastic debris in the open ocean. Proc. Natl Acad. Sci. USA 111, 10239–10244 (2014).
Google Scholar 
51.Mattsson, K., Björkroth, F., Karlsson, T. & Hassellöv, M. Nanofragmentation of expanded polystyrene under simulated environmental weathering (thermooxidative degradation and hydrodynamic turbulence). Front. Mar. Sci., 7, 1–9 (2021). This paper demonstrates log linear particle size distributions extending to the nanoparticle scale.
Google Scholar 
52.Kaandorp, M. L. A., Dijkstra, H. A. & van Sebille, E. Modelling size distributions of marine plastics under the influence of continuous cascading fragmentation. Environ. Res. Lett. 16, 054075 (2021).
Google Scholar 
53.Koelmans, A. A., Besseling, E. & Shim, W. J. Nanoplastics in the aquatic environment. Critical review. In Marine Anthropogenic Litter (eds Bergmann, M., Gutow, L. & Klages, M.) 325–340 (Springer, 2015).54.Koelmans, A. A. et al. Microplastics in freshwaters and drinking water: critical review and assessment of data quality. Water Res. 155, 410–422 (2019).CAS 

Google Scholar 
55.Chamas, A. et al. Degradation rates of plastics in the environment. ACS Sust. Chem. Engin. 8, 3494–3511 (2020). This paper provides a rare estimate of degradation rates for plastic items in the environment.CAS 

Google Scholar 
56.Unice, K. M. et al. Characterizing export of land-based microplastics to the estuary — Part II: Sensitivity analysis of an integrated geospatial microplastic transport modeling assessment of tire and road wear particles. Sci. Total. Environ. 646, 1650–1659 (2019).CAS 

Google Scholar 
57.Buffle, J. & van Leeuwen, H. P. Environmental Particles Vol. 1 76 (CRC Press, 1992).58.Chamley, H., Clay formation through weathering. In Clay Sedimentology (Springer, 1989).59.Blott, S. J. & Pye, K. Particle size scales and classification of sediment types based on particle size distributions: review and recommended procedures. Sedimentology 59, 2071–2096 (2012).
Google Scholar 
60.Boyd, C. E. Suspended solids, color, turbidity, and light. In Water Quality 119–133 (Springer, 2020).61.Konrad, K. et al. Chemical composition and complex refractive index of Saharan mineral dust at Izaña, Tenerife (Spain) derived by electron microscopy. Atmos. Env. 41, 8058–8074 (2007).
Google Scholar 
62.Mahowald, N. et al. The size distribution of desert dust aerosols and its impact on the Earth system. Aeolian Res. 15, 53–71 (2014).
Google Scholar 
63.De Wit, C. T. & Arens, P. L. Moisture content and density of some clay minerals and some remarks on the hydration pattern of clay. Trans. Int. Congr. Soil Science 2, 59–62 (1951).
Google Scholar 
64.Utembe, W., Potgieter, K., Stefaniak, A. B. & Gulumian, M. Dissolution and biodurability: important parameters needed for risk assessment of nanomaterials. Part. Fiber Toxicol. 12, 11 (2015).
Google Scholar 
65.Köhler, S. J., Bosbach, D. & Oelkers, E. H. Do clay mineral dissolution rates reach steady state? Geochim. Cosmochim. Acta 69, 1997–2006 (2005).
Google Scholar 
66.Torrey, M. L. S. T. Chemistry of Lake Michigan (Argonne National Laboratory, 1976).67.Prestigiacomo, A. R. et al. Turbidity and suspended solids levels and loads in a sediment enriched stream: implications for impacted lotic and lentic ecosystems. Lake Res. Manag. 23, 231–244 (2007).
Google Scholar 
68.Baran, A. et al. The influence of the quantity and quality of sediment organic matter on the potential mobility and toxicity of trace elements in bottom sediment. Environ. Geochem. Health 41, 2893–2910 (2019).CAS 

Google Scholar 
69.Schwarzenbach, R. P., Gschwend, P. M. & Imboden, D. M. Environmental Organic Chemistr 3rd edn 1024 (Wiley, 2016).70.Van Valkenburg, S. D., Jones, J. K. & Heinle, D. R. A comparison by size class and volume of detritus versus phytoplankton in Chesapeake Bay. Estuar. Coast. Mar. Sci. 6, 569–582 (1978).
Google Scholar 
71.Hamilton, S. K., Sippel, S. J. & Bunn, S. E. Separation of algae from detritus for stable isotope or ecological stoichiometry studies using density fractionation in colloidal silica. Limnol. Oceanogr. Methods 3, 149–157 (2005).CAS 

Google Scholar 
72.Zimmer, M. Detritus. Encyclopedia of Ecology 903–911 (Elsevier, 2008).73.Zhao, H.-C., Wang, S.-R., Jiao, L.-X., Yang, S.-W. & Cui, C.-N. Characteristics of composition and spatial distribution of organic matter in the sediment of Erhai Lake. Res. Environ. Sci. 26, 243–249 (2013).CAS 

Google Scholar 
74.Duan, H., Feng, L., Ma, R., Zhang, Y. & Loiselle, S. A. Variability of particulate organic carbon in inland waters observed from MODIS Aqua imagery. Environ. Res. Lett. 9, 084011 (2014).CAS 

Google Scholar 
75.Suaria, G. et al. Microfibers in oceanic surface waters: a global characterization. Sci. Adv. 6, eaay8493 (2020). This paper identifies the relative proportion of microplastic fibres in the oceans.
Google Scholar 
76.Le Guen, C. et al. Microplastic study reveals the presence of natural and synthetic fibres in the diet of King penguins (Aptenodytes patagonicus) foraging from South Georgia. Environ. Intern. 134, 105303 (2020).
Google Scholar 
77.Stanton, T., Johnson, M., Nathanail, P., MacNaughtan, W. & Gomes, R. L. Sci. Total. Environ. 666, 377–389 (2019).CAS 

Google Scholar 
78.Comnea-Stancu, L. R., Wieland, H., Ramer, G., Schwaighofer, A. & Lendl, B. On the identification of rayon/viscose as a major fraction of microplastics in the marine environment: discrimination between natural and manmade cellulosic fibers using Fourier transform infrared spectroscopy. Appl. Spectrosc. 71, 939–950 (2017).CAS 

Google Scholar 
79.Seiler, W. & Crutzen, P. J. Estimates of gross and net fluxes of carbon between the biosphere and the atmosphere from biomass burning. Clim. Change 2, 207–247 (1980).CAS 

Google Scholar 
80.Cornelissen, G. et al. Critical review. Extensive sorption of organic compounds to black carbon, coal, and kerogen in sediments and soils: mechanisms and consequences for distribution, bioaccumulation and biodegradation. Environ. Sci. Technol. 39, 6881–6895 (2005).CAS 

Google Scholar 
81.Jonker, M. T. O., Hawthorne, S. B. & Koelmans, A. A. Extremely slow desorption and limited bioaccumulation of BC-associated PAHs. ACS Div. Environ. Chem. 45, 381–384 (2005).CAS 

Google Scholar 
82.Shrestha, G., Traina, S. J., Swanson & C., W. Black carbons properties and role in the environment: a comprehensive review. Sustainability 2, 294–320 (2010).CAS 

Google Scholar 
83.Bisiaux, M. M. et al. Stormwater and fire as sources of black carbon nanoparticles to Lake Tahoe. Environ. Sci. Technol. 45, 2065–2071 (2011). This paper identifies black carbon abundance in surface waters.CAS 

Google Scholar 
84.World Health Organization. Health effects of black carbon. (WHO, 2012).85.Jonker, M. T. O. & Koelmans, A. A. Sorption of polycyclic aromatic hydrocarbons and polychlorinated biphenyls to soot and soot-like materials in the aqueous environment. Mechanistic considerations. Environ. Sci. Technol. 36, 3725–3734 (2002).CAS 

Google Scholar 
86.Liu, H. et al. Mixing characteristics of refractory black carbon aerosols at an urban site in Beijing. Atmos. Chem. Phys. 20, 5771–5785 (2020).CAS 

Google Scholar 
87.Ouf, F.-X. et al. True density of soot particles: a comparison of results highlighting the influence of the organic contents. J. Aerosol Sci. 134, 1–13 (2019).CAS 

Google Scholar 
88.Wu, Y. et al. A study of the morphology and effective density of externally mixed black carbon aerosols in ambient air using a size-resolved single-particle soot photometer (SP2). Atmos. Meas. Tech. 12, 4347–4359 (2019).CAS 

Google Scholar 
89.Kuzyakov, Y., Subbotina, I., Chen, H., Bogomolova, I. & Xu, X. Black carbon decomposition and incorporation into soil microbial biomass estimated by 14C labeling. Soil. Biol. Biochem. 41, 210–219 (2009).CAS 

Google Scholar 
90.Middelburg, J. J., Nieuwenhuize, J. & Van Breugel, P. Black carbon in marine sediments. Mar. Chem. 65, 245–252 (1999).CAS 

Google Scholar 
91.Murr, L. E., Bang, J. J., Esquivel, E. V., Guerrero, P. A. & Lopez, D. A. Carbon nanotubes, nanocrystal forms and complex nanoparticle aggregates in common fuel gas combustion streams. J. Nanopart. Res. 6, 241–251 (2004).CAS 

Google Scholar 
92.Koelmans, A. A., Nowack, B. & Wiesner, M. Comparison of manufactured and black carbon nanoparticle concentrations in aquatic sediments. Environ. Pollut. 157, 1110–1116 (2009).CAS 

Google Scholar 
93.Dickens, A. F., Gelinas, Y., Masiello, C. A., Wakeham, S. & Hedges, J. I. Reburial of fossil organic carbon in marine sediments. Nature 427, 336–339 (2004).CAS 

Google Scholar 
94.Kharbush, J. J. et al. Particulate organic carbon deconstructed: molecular and chemical composition of particulate organic carbon in the ocean. Front. Mar. Sci. 7, 518 (2020).
Google Scholar 
95.Redondo-Hasselerharm, P. E. Effect assessment of nano- and microplastics in freshwater ecosystems. Thesis, Wageningen Univ. (2020).96.Allen, S. et al. Atmospheric transport and deposition of microplastics in a remote mountain catchment. Nat. Geosci. 12, 339–344 (2019).CAS 

Google Scholar 
97.Evangeliou, N. et al. Atmospheric transport is a major pathway of microplastics to remote regions. Nat. Commun. 11, 3381 (2020).CAS 

Google Scholar 
98.Velzeboer, I., Kwadijk, C. J. A. F. & Koelmans, A. A. Strong sorption of PCBs to nanoplastics, microplastics, carbon nanotubes and fullerenes. Environ. Sci. Technol. 48, 4869–4876 (2014).CAS 

Google Scholar 
99.Beckingham, B. & Ghosh, U. Differential bioavailability of polychlorinated biphenyls associated with environmental particles: microplastic in comparison to wood, coal and biochar. Environ. Pollut. 220, 150–158 (2017).CAS 

Google Scholar 
100.Liping, L. et al. Mechanism of and relation between the sorption and desorption of nonylphenol on black carbon-inclusive sediment. Environ. Pollut. 190, 101–108 (2014).
Google Scholar 
101.Voparil, I. M. et al. Digestive bioavailability to a deposit feeder (Arenicola marina) of polycyclic aromatic hydrocarbons associated with anthropogenic particles. Environ. Toxicol. Chem. 23, 2618–2626 (2004).CAS 

Google Scholar 
102.Birdwell, J., Cook, R. L. & Thibodeaux, L. J. Desorption kinetics of hydrophobic organic chemicals from sediment to water: a review of data and models. Environ. Toxicol. Chem. 26, 424–434 (2007).CAS 

Google Scholar 
103.Koelmans, A. A., Besseling, E. & Foekema, E. M. Leaching of plastic additives to marine organisms. Environ. Pollut. 187, 49–54 (2014).CAS 

Google Scholar 
104.Bundschuh, M. et al. Nanoparticles in the environment: where do we come from, where do we go to? Environ. Sci. Eur. 30, 6 (2018).
Google Scholar 
105.Peijnenburg, W. J. G. M. et al. A review of the properties and processes determining the fate of engineered nanomaterials in the aquatic environment. Crit. Rev. Environ. Sci. Technol. 45, 2084–2134 (2015).CAS 

Google Scholar 
106.Gigault, J. et al. Nanoplastics are neither microplastics nor engineered nanoparticles. Nat. Nanotechnol. 16, 501–507 (2021).CAS 

Google Scholar 
107.Ter Halle, A. et al. Nanoplastic in the North Atlantic Subtropical Gyre. Environ. Sci. Technol. 51, 13689–13697 (2017).
Google Scholar 
108.Sengul, A. B. & Asmatulu, E. Toxicity of metal and metal oxide nanoparticles: a review. Environ. Chem. Lett. 18, 1659–1683 (2020).CAS 

Google Scholar 
109.Botterell, Z. L. R. et al. Bioavailability and effects of microplastics on marine zooplankton: a review. Environ. Pollut. 245, 98–110 (2019).CAS 

Google Scholar 
110.Ribeiro, F., O’Brien, J. W., Galloway, T. & Thomas, K. V. Accumulation and fate of nano- and micro-plastics and associated contaminants in organisms. TrAC 111, 139–147 (2019).CAS 

Google Scholar 
111.da Costa Araújo, A. P. et al. How much are microplastics harmful to the health of amphibians? A study with pristine polyethylene microplastics and Physalaemus cuvieri. J. Hazard. Mater. 382, 121066 (2020).
Google Scholar 
112.Jovanović, B. Ingestion of microplastics by fish and its potential consequences from a physical perspective. Integr. Environ. Assess. Manag. 13, 510–515 (2017).
Google Scholar 
113.Windsor, F. M., Tilley, R. M., Tyler, C. R. & Ormerod, S. J. Microplastic ingestion by riverine macroinvertebrates. Sci. Total. Environ. 646, 68–74 (2018).
Google Scholar 
114.Hu, L., Chernick, M., Hinton, D. E. & Shi, H. Microplastics in small waterbodies and tadpoles from Yangtze River Delta, China. Environ. Sci. Technol. 52, 8885–8893 (2018).CAS 

Google Scholar 
115.McNeish, R. E. et al. Microplastic in riverine fish is connected to species traits. Sci. Rep. 8, 11639 (2018).CAS 

Google Scholar 
116.Duncan, E. M. et al. Microplastic ingestion ubiquitous in marine turtles. Glob. Chang. Biol. 25, 744–752 (2019).
Google Scholar 
117.Kühn, S., Bravo Rebolledo, E. L. & Van Franeker, J. A. Deleterious effects of litter on marine life. In Marine Anthropogenic Litter (eds Bergmann, M., Gutow, L. & Klages, M.) 75–116 (Springer International Publishing, 2015).118.Nelms, S. E. et al. Microplastics in marine mammals stranded around the British coast: ubiquitous but transitory? Sci. Rep. 9, 1–9 (2019).CAS 

Google Scholar 
119.O’Connor, J. D. et al. Microplastics in freshwater biota: a critical review of isolation, characterization, and assessment methods. Glob. Challen. 4, 1800118 (2019).
Google Scholar 
120.Vroom, R. J. E., Koelmans, A. A., Besseling, E. & Halsband, C. Aging of microplastics promotes their ingestion by marine zooplankton. Environ. Pollut. 231, 987–996 (2017).CAS 

Google Scholar 
121.Bour, A., Haarr, A., Keiter, S. & Hylland, K. Environmentally relevant microplastic exposure affects sediment-dwelling bivalves. Environ. Pollut. 236, 652–660 (2018).CAS 

Google Scholar 
122.Kaposi, K. L., Mos, B., Kelaher, B. P. & Dworjanyn, S. A. Ingestion of microplastic has limited impact on a marine larva. Environ. Sci. Technol. 48, 1638–1645 (2014).CAS 

Google Scholar 
123.Lu, Y. et al. Uptake and accumulation of polystyrene microplastics in zebrafish (Danio rerio) and toxic effects in liver. Environ. Sci. Technol. 50, 4054–4060 (2016).CAS 

Google Scholar 
124.Ribeiro, F. et al. Microplastics effects in Scrobicularia plana. Mar. Pollut. Bull. 122, 379–391 (2017).CAS 

Google Scholar 
125.Von Moos, N., Burkhardt-Holm, P. & Köhler, A. Uptake and effects of microplastics on cells and tissue of the blue mussel Mytilus edulis L. after an experimental exposure. Environ. Sci. Technol. 46, 11327–11335 (2012).
Google Scholar 
126.Browne, M. A., Dissanayake, A., Galloway, T. S., Lowe, D. M. & Thompson, R. C. Ingested microscopic plastic translocates to the circulatory system of the mussel, Mytilus edulis (L.). Environ. Sci. Technol. 42, 5026–5031 (2008).CAS 

Google Scholar 
127.Dawson, A. L. et al. Turning microplastics into nanoplastics through digestive fragmentation by Antarctic krill. Nat. Commun. 9, 1001 (2018).
Google Scholar 
128.Zhang, C., Chen, X., Wang, J. & Tan, L. Toxic effects of microplastic on marine microalgae Skeletonema costatum: interactions between microplastic and algae. Environ. Pollut. 220, 1282–1288 (2017).CAS 

Google Scholar 
129.Mateos-Cárdenas, A. et al. Polyethylene microplastics adhere to Lemna minor (L.), yet have no effects on plant growth or feeding by Gammarus duebeni (Lillj.). Sci. Total. Environ. 689, 413–421 (2019).
Google Scholar 
130.Murphy, F. & Quinn, B. The effects of microplastic on freshwater Hydra attenuata feeding, morphology and reproduction. Environ. Pollut. 234, 487–494 (2018).CAS 

Google Scholar 
131.Cole, M. et al. Microplastic ingestion by zooplankton. Environ. Sci. Technol. 47, 6646–6655 (2013).CAS 

Google Scholar 
132.Green, D. S., Boots, B., O’Connor, N. E. & Thompson, R. Microplastics affect the ecological functioning of an important biogenic habitat. Environ. Sci. Technol. 51, 68–77 (2017).CAS 

Google Scholar 
133.Senga Green, D. Effects of microplastics on European flat oysters, Ostrea edulis and their associated benthic communities. Environ. Pollut. 216, 95–103 (2016).
Google Scholar 
134.Ziajahromi, S., Kumar, A., Neale, P. A. & Leusch, F. D. L. Environmentally relevant concentrations of polyethylene microplastics negatively impact the survival, growth and emergence of sediment-dwelling invertebrates. Environ. Pollut. 236, 425–431 (2018).CAS 

Google Scholar 
135.Ogonowski, M., Schür, C., Jarsén, Å. & Gorokhova, E. The effects of natural and anthropogenic microparticles on individual fitness in daphnia magna. PLoS ONE 11, e0155063 (2016). This paper systematically addresses the differences between the biological effects of microplastic and natural particles.
Google Scholar 
136.Mazurais, D. et al. Evaluation of the impact of polyethylene microbeads ingestion in European sea bass (Dicentrarchus labrax) larvae. Mar. Environ. Res. 112, 78–85 (2015).CAS 

Google Scholar 
137.Lee, K.-W., Shim, W. J., Kwon, O. Y. & Kang, J.-H. Size-dependent effects of micro polystyrene particles in the marine copepod Tigriopus japonicus. Env. Sci. Technol. 47, 11278–11283 (2013).CAS 

Google Scholar 
138.Au, S. Y., Bruce, T. F., Bridges, W. C. & Klaine, S. J. Responses of Hyalella azteca to acute and chronic microplastic exposures. Environ. Toxicol. Chem. 34, 2564–2572 (2015).CAS 

Google Scholar 
139.Cole, M., Lindeque, P., Fileman, E., Halsband, C. & Galloway, T. S. The impact of polystyrene microplastics on feeding, function and fecundity in the marine copepod Calanus helgolandicus. Environ. Sci. Technol. 49, 1130–1137 (2015).CAS 

Google Scholar 
140.Sussarellu, R. et al. Oyster reproduction is affected by exposure to polystyrene microplastics. Proc. Natl Acad. Sci. USA 113, 2430–2435 (2016).CAS 

Google Scholar 
141.Jeong, C. B. et al. Microplastic size-dependent toxicity, oxidative stress induction, and p-JNK and p-p38 activation in the monogonont rotifer (Brachionus koreanus). Environ. Sci. Technol. 50, 8849–8857 (2016).CAS 

Google Scholar 
142.Blarer, P. & Burkhardt-Holm, P. Microplastics affect assimilation efficiency in the freshwater amphipod Gammarus fossarum. Environ. Sci. Pollut. Res. 23, 23522–23532 (2016).CAS 

Google Scholar 
143.Wright, S. L., Rowe, D., Thompson, R. C. & Galloway, T. S. Microplastic ingestion decreases energy reserves in marine worms. Curr. Biol. 23, R1031–R1033 (2013).CAS 

Google Scholar 
144.Straub, S., Hirsch, P. E. & Burkhardt-Holm, P. Biodegradable and petroleum-based microplastics do not differ in their ingestion and excretion but in their biological effects in a freshwater invertebrate Gammarus fossarum. Int. J. Environ. Res. Public Health 14, 774 (2017).
Google Scholar 
145.Green, D. S., Boots, B., Sigwart, J., Jiang, S. & Rocha, C. Effects of conventional and biodegradable microplastics on a marine ecosystem engineer (Arenicola marina) and sediment nutrient cycling. Environ. Pollut. 208, 426–434 (2016).CAS 

Google Scholar 
146.Ziajahromi, S., Kumar, A., Neale, P. A. & Leusch, F. D. L. Impact of microplastic beads and fibers on waterflea (Ceriodaphnia dubia) survival, growth, and reproduction: implications of single and mixture exposures. Environ. Sci. Technol. 51, 13397–13406 (2017).CAS 

Google Scholar 
147.Nobre, C. R. et al. Assessment of microplastic toxicity to embryonic development of the sea urchin Lytechinus variegatus (Echinodermata: Echinoidea). Mar. Pollut. Bull. 92, 99–104 (2015).CAS 

Google Scholar 
148.Rehse, S., Kloas, W. & Zarfl, C. Short-term exposure with high concentrations of pristine microplastic particles leads to immobilisation of Daphnia magna. Chemosphere 153, 91–99 (2016).CAS 

Google Scholar 
149.Gambardella, C. et al. Effects of polystyrene microbeads in marine planktonic crustaceans. Ecotoxicol. Environ. Saf. 145, 250–257 (2017).CAS 

Google Scholar 
150.Watts, A. J. R. et al. Effect of microplastic on the gills of the shore crab Carcinus maenas. Environ. Sci. Technol. 50, 5364–5369 (2016).CAS 

Google Scholar 
151.Espinosa, C., Cuesta, A. & Esteban, M. Á. Effects of dietary polyvinylchloride microparticles on general health, immune status and expression of several genes related to stress in gilthead seabream (Sparus aurata L.). Fish. Shellfish. Immunol. 68, 251–259 (2017).CAS 

Google Scholar 
152.Jin, Y., Lu, L., Tu, W., Luo, T. & Fu, Z. Impacts of polystyrene microplastic on the gut barrier, microbiota and metabolism of mice. Sci. Total. Environ. 649, 308–317 (2019).CAS 

Google Scholar 
153.Jin, Y. et al. Polystyrene microplastics induce microbiota dysbiosis and inflammation in the gut of adult zebrafish. Environ. Pollut. 235, 322–329 (2018).CAS 

Google Scholar 
154.Bucci, K., Tulio, M. & Rochman, C. M. What is known and unknown about the effects of plastic pollution: a meta-analysis and systematic review. Ecol. Appl. 30, e02044 (2020). This paper reviews the evidence for effects of plastic pollution across endpoints, organisms and levels of biological organization.CAS 

Google Scholar 
155.Kjelland, M. E., Woodley, C. M., Swannack, T. M. & Smith, D. L. A review of the potential effects of suspended sediment on fishes: potential dredging-related physiological, behavioral, and transgenerational implications. Environ. Syst. Decis. 35, 334–350 (2015).
Google Scholar 
156.Michel, C., Herzog, S., de Capitani, C., Burkhardt-Holm, P. & Pietsch, C. Natural mineral particles are cytotoxic to rainbow trout gill epithelial cells in vitro. PLoS ONE 9, e100856 (2014).
Google Scholar 
157.Gordon, A. K. & Palmer, C. G. Defining an exposure-response relationship for suspended kaolin clay particulates and aquatic organisms: work toward defining a water quality guideline for suspended solids. Environ. Toxicol. Chem. 34, 907–912 (2015).CAS 

Google Scholar 
158.Lu, C., Kania, P. W. & Buchmann, K. Particle effects on fish gills: an immunogenetic approach for rainbow trout and zebrafish. Aquaculture 484, 98–104 (2018).CAS 

Google Scholar 
159.Ogonowski, M., Gerdes, Z. & Gorokhova, E. What we know and what we think we know about microplastic effects — a critical perspective. Curr. Opin. Environ. Sci. Health 1, 41–46 (2018).
Google Scholar 
160.Albarano, L. et al. Comparison of in situ sediment remediation amendments: risk perspectives from species sensitivity distribution. Environ. Pollut. 272, 115995 (2021).CAS 

Google Scholar 
161.Newcombe, C. P. & Macdonald, D. D. Effects of suspended sediments on aquatic ecosystems. North. Am. J. Fish. Manag. 11, 72–82 (1991).
Google Scholar 
162.Yap, V. H. et al. A comparison with natural particles reveals a small specific effect of PVC microplastics on mussel performance. Mar. Pollut. Bull. 160, 111703 (2020).CAS 

Google Scholar 
163.Schür, C., Zipp, S., Thalau, T. & Wagner, M. Microplastics but not natural particles induce multigenerational effects in Daphnia magna. Environ. Pollut. 260, 113904 (2020).
Google Scholar 
164.Gerdes, Z., Hermann, M., Ogonowski, M. & Gorokhova, E. A novel method for assessing microplastic effect in suspension through mixing test and reference materials. Sci. Rep. 9, 1–9 (2019).CAS 

Google Scholar 
165.Niranjan, R. & Thakur, A. K. The toxicological mechanisms of environmental soot (black carbon) and carbon black: focus on oxidative stress and inflammatory pathways. Front. Immunol. 8, 763 (2017).
Google Scholar 
166.Tsuji, J. S. et al. Research strategies for safety evaluation of nanomaterials. Part IV: Risk assessment of nanoparticles. Toxicol. Sci. 89, 42–50 (2006).CAS 

Google Scholar 
167.Schwarze, P. E. et al. Importance of size and composition of particles for effects on cells in vitro. Inhal. Toxicol. 19, 17–22 (2007).CAS 

Google Scholar 
168.Schmid, O. & Stoeger, T. Surface area is the biologically most effective dose metric for acute nanoparticle toxicity in the lung. J. Aerosol Sci. 99, 133–143 (2016). This paper identifies the toxicologically relevant dose metric for particle effects.CAS 

Google Scholar 
169.Fubini, B. Surface reactivity in the pathogenic response to particulates. Environ. Health Perspect. 105, 1013–1020 (1997).
Google Scholar 
170.Poland, C. A., Duffin, R. & Donaldson, K. High aspect ratio nanoparticles and the fibre pathogenicity paradigm. In Nanotoxicity Vivo and In Vitro Models to Health Risks 61–80 (John Wiley and Sons, 2009).171.Gualtieri, A. F. Bridging the gap between toxicity and carcinogenicity of mineral fibres by connecting the fibre crystal-chemical and physical parameters to the key characteristics of cancer. Curr. Res. Toxicol. 2, 42–52 (2021).
Google Scholar 
172.Shao, X. R. et al. Independent effect of polymeric nanoparticle zeta potential/surface charge, on their cytotoxicity and affinity to cells. Cell Prolif. 48, 465–474 (2015).CAS 

Google Scholar 
173.Motskin, M. et al. Hydroxyapatite nano and microparticles: correlation of particle properties with cytotoxicity and biostability. Biomaterials 30, 3307–3317 (2009).CAS 

Google Scholar 
174.Cox, K. D. et al. Human consumption of microplastics. Environ. Sci. Technol. 53, 7068–7074 (2019).CAS 

Google Scholar 
175.Zhang, Q. et al. A review of microplastics in table salt, drinking water, and air: direct human exposure. Environ. Sci. Technol. 54, 3740–3751 (2020).CAS 

Google Scholar 
176.Everaert, G. et al. Risk assessment of microplastics in the ocean: modelling approach and first conclusions. Environ. Pollut. 242, 1930–1938 (2018).CAS 

Google Scholar 
177.Everaert, G. et al. Risks of floating microplastic in the global ocean. Environ. Pollut. 267, 115499 (2020).CAS 

Google Scholar 
178.Zhang, X., Leng, Y., Liu, X., Huang, K. & Wang, J. Microplastics’ pollution and risk assessment in an urban river: a case study in the Yongjiang River, Nanning City, South China. Exposure Health 12, 141–151 (2020).CAS 

Google Scholar 
179.Skåre, J. U. et al. Microplastics, occurrence, levels and implications for environment and human health related to food. Opinion of the steering committee of the Norwegian Scientific Committee for Food and Environment (VKM, 2019).180.Adam, V., von Wyl, A. & Nowack, B. Probabilistic environmental risk assessment of microplastics in marine habitats. Aq. Toxicol. 230, 105689 (2021).CAS 

Google Scholar 
181.Jung, J.-W. et al. Ecological risk assessment of microplastics in coastal, shelf, and deep sea waters with a consideration of environmentally relevant size and shape. Environ. Pollut. 270, 116217 (2021).CAS 

Google Scholar 
182.Posthuma, L., Suter, G. W. & Traas, T. P. Species Sensitivity Distributions In Ecotoxicology (Lewis, 2002).183.Gouin, T. et al. Toward the development and application of an environmental risk assessment framework for microplastic. Environ. Toxicol. Chem. 38, 2087–2100 (2019).CAS 

Google Scholar 
184.Kong, X. & Koelmans, A. A. Effects of microplastic on shallow lake food webs. Environ. Sci. Technol. 53, 13822–13831 (2019).CAS 

Google Scholar 
185.Zimmermann, L., Göttlich, S., Oehlmann, J., Wagner, M. & Völker, C. What are the drivers of microplastic toxicity? Comparing the toxicity of plastic chemicals and particles to Daphnia magna. Environ. Pollut. 267, 115392 (2020).CAS 

Google Scholar 
186.Tian, Z. et al. A ubiquitous tire rubber-derived chemical induces acute mortality in coho salmon. Science 371, 185–189 (2021).CAS 

Google Scholar 
187.Bakir, A., O’Connor, I. A., Rowland, S. J., Hendriks, A. J. & Thompson, R. C. Relative importance of microplastics as a pathway for the transfer of hydrophobic organic chemicals to marine life. Environ. Pollut. 219, 56–65 (2016).CAS 

Google Scholar 
188.Capolupo, M., Sørensen, L., Jayasena, K., Booth, A. M. & Fabbri, E. Chemical composition and ecotoxicity of plastic and car tire rubber leachates to aquatic organisms. Water Res. 169, 115270 (2020).CAS 

Google Scholar 
189.Zimmermann, L. et al. Plastic products leach chemicals that induce in vitro toxicity under realistic use conditions. Environ. Sci. Technol. 55, 11814–11823 (2021).CAS 

Google Scholar 
190.Bucci, K. & Rochman, C. M. A proposed framework for microplastics risk assessment [abstract 07.05.02]. Society of Environmental Toxicology and Chemistry North America 42nd Annual Meeting – SETAC SciCon4 https://scicon4.setac.org/wp-content/uploads/2021/11/SciCon4-abstract-book.pdf (2021).191.Primpke, S., Lorenz, C., Rascher-Friesenhausen, R. & Gerdts, G. An automated approach for microplastics analysis using focal plane array (FPA) FTIR microscopy and image analysis. Anal. Methods 9, 1499–1511 (2017).CAS 

Google Scholar 
192.Rauchschwalbe, M.-T., Fueser, H., Traunspurger, W. & Höss, S. Bacterial consumption by nematodes is disturbed by the presence of polystyrene beads: the roles of food dilution and pharyngeal pumping. Environ. Pollut. 273, 116471 (2021).CAS 

Google Scholar 
193.Donaldson, K. & Seaton, A. A short history of the toxicology of inhaled particles. Part. Fibre Toxicol. 9, 13 (2012).
Google Scholar 
194.Primpke, S., Dias, A. P. & Gerdts, G. Automated identification and quantification of microfibers and microplastics. Anal. Methods 11, 2138–2147 (2019).CAS 

Google Scholar  More

Our results show that differences in environmental conditions between inland and coastal sinkholes, caused mainly by the inflow of seawater in the latter, influence the microbial community structure of their sediments. Furthermore, the microbial community structure also varied within the sinkholes and according to the sediment zone sampled, suggesting that a connection between the atmosphere in the outermost location of the sediments and sunlight creates an environment distinct from that found in deeper caves. Together with the different environmental factors that were measured (in situ physicochemical composition of water and sediment) these characteristics could drive niche-specific microbial community structures associated with the sediment zones. Additionally, beta-diversity analysis showed separate clustering of the sediment microbial communities from the coastal and inland sinkholes, and of the WM zone from cavern and cave zones at both sinkholes. Microbial community structure associated with karst environments have shown to be significantly influenced by environmental factors as seen in the Bahamian blue holes19, a coastal sinkhole13, a Floridan anchialine sinkhole20, and sediments from Chinese karst caves21.Microbial communities from karst sediments can be limited by nutrients such as carbon, phosphorus, and nitrogen21, therefore, influencing their structure. Differences in the microbial community composition associated with multiple environmental factors (moisture, type of niche, nitrogen) were also reported in karst cave sediments from China21. Previous studies had shown there was no effect on the alpha diversity of water column assemblages in the Yucatan groundwater associated with the type of sinkhole (inland or coastal)6. However, this observation may be limited due to the low sample number used in the study6. For other karst sinkholes, the microbial community dynamics differ between the water column and the sediments6,22,23,24.The karst caves and sinkholes of the underground river in Yucatan are characterized by low phosphorus concentrations and high levels of nitrate, mostly related to Anthropocentric activities (urban developments, farms and agriculture)3. The inland sinkhole at Noh Mozón showed the highest concentration of nitrate detected in the study and, not surprisingly, the area is surrounded by agricultural fields. The presence of organic matter in the sinkholes from the Yucatan peninsula are highly dependent on the connection between the cave systems, on the levels of exposure to light, and on their morphology3. High concentrations of organic carbon (661 ± 132 μM) and methane (6466 ± 659 nM) have been reported in the top layer of the water masses in coastal sinkholes before13. In this study, the highest concentration of organic carbon was observed in the sediments from the coastal sinkhole, likely originating from the surrounding vegetation and from seawater intrusion. We hypothesize that the differences in nutrients found at these two types of sinkholes influence the structure of their microbial communities.Other environmental factors such as pH and dissolved oxygen (DO), may also contribute significantly to the composition and structure of microbial communities, as seen in freshwater lake sediments22. Davis and Garey20 reported distinct microbial communities with unique functions for each water layer from an anchialine sinkhole from the Florida karst aquifer and suggested that this occurred as a result of the influence of the hydrochemistry, including differences in the concentration carbon and other nutrients from the environment20. Analyses of the sinkhole caves from the Yucatan underwater river support observations that physical and chemical parameters create distinct ecological niches which host unique microbes, as a high abundance of exclusive (not shared) ASVs were observed in the three sediment zones at both locations.The taxonomic diversity from the coastal and inland sinkholes included Chloroflexi, Crenarchaeota, Desulfobacterota, Proteobacteria, Nitrospirota, Bacteroidota, and Firmicutes as the most abundant phyla in the sediment samples, however, there were differences in the relative abundance associated with the type of sinkhole and sediment zone. Some of these phyla (Chloroflexi, Proteobacteria, and Bacteroidetes) have been reported in sediments from freshwater karst sinkholes from Lake Huron25 in water and sediments from other sinkholes in the Yucatan peninsula6, and in the karst caves bacteriome from southwest China21. A study that included coastal marine sediments from two sites in the Yucatan peninsula, showed high abundances of Spirochaeta, Desulfococcus, Clostridium, Psychrobacter26, four genera that were abundant in the coastal sinkhole. However, Desulfococcus, Synechococcus were also abundant in the inland sinkhole. Of the most abundant genera reported for sediments from different marine environments in the Yucatan coast23, Acinetobacter, Desulfotignum, Desulfovibrio, Pseudomonas, Sedimenticola, and Sulfurimonas were also present in the coastal sinkhole while only Pseudomonas and Sedimenticola were also present in the inland sinkhole23. The high number of families shared between the coastal sinkhole and marine sediments from the Yucatan coast, together with the salinity levels registered at the bottom layer of the water column in the coastal sinkhole, suggest an interconnection between these two environments which shapes the microbial communities present in the sediments of caverns and caves of this sinkhole. The genus Nitrospira was abundant in the WM from the coastal sinkhole and in all sediment zones from the inland sinkhole. This genus has been reported as one of the most abundant in the surface of speleothems from El Zapote coastal sinkhole2, and is considered a complete ammonia oxidizer (comammox), meaning it converts ammonia to nitrate through nitrite. A negative correlation between abundance of this genus and salinity has been reported before, which could explain the low concentration of Nitrospira in the cavern and cave from the coastal sinkhole, where the highest salinity was observed27. Connectivity between coastal sinkholes and the ocean, as well as the terrestrial input of soil organic matter (OM) has been reported for the underground karst aquifer in the Yucatan peninsula13. As in other sediments, degradation of OM is carried out by several MFGs including acetogenic bacteria, methanogens, and sulfate reducers13,20,28. When these MFGs were analyzed in coastal and inland sinkholes, differences in their relative abundances were clearly marked by the type of sinkhole and by the sediment zone analyzed, supporting the hypothesis that environmental differences drive microbial community distributions in these niches. The high abundance of sulfate-reducing bacteria (SRB) in the three sediment zones from the coastal sinkhole suggests that sulfate reduction is a predominant function. SRB degrade organic matter using sulfate with sulfide as waste or end-product19,30, originating hydrogen sulfide (H2S)29, which could explain the low concentration of sulfate the hydrogen sulfide (H2S) cloud observed and previously reported in the WM zone of El Zapote coastal sinkhole30. In this study, high levels of sulfate (SO−4) were measured in the water samples from the cavern and cave zones from El Zapote sinkhole, which could be associated with sulfate-rich deposits, such as gypsum beds, which have been reported in other sinkholes from the Yucatan peninsula (up to 2400 mg/L of sulfate concentration)3. However, we do not disregard other possible sources of sulfate, associated with seawater intrusion or as a product of sulfide or sulfur oxidation29,31 by sulfur-oxidizing bacteria detected in this study.The inland sinkhole had a low concentration of sulfate and low abundances of SRB. The high abundance of methanogenic bacteria in the WM zone from the coastal sinkhole detected in the MFG analysis supports the previous hypothesis of acetoclastic methanogenesis due to high inputs of organic matter13. Methylotrophic bacteria were most abundant at the inland sinkhole in the WM zone, suggesting the presence of methyl compounds, such as methane or methanol which can be used as a source of carbon and energy32. High methane concentrations have been quantified in shallow water masses from the Yucatan aquifer system13, consistent with observations from this study. ‘Candidatus Methylomirabilis’ was identified in the sediment of the WM zone from Noh-Mozón and has been previously described as being able to perform nitrite-dependent anaerobic methane oxidation, using methane as electron donor and nitrate and nitrite as electron acceptors33, which would be possible in these sediments considering the low levels of oxygen (average of 2.3 mg/L) detected in the water column above them and assuming this would lead to lower levels of oxygen in the sediments. We hypothesize that bacteria from this genus could be using the nitrite produced by ammonia oxidizing bacteria and archaea observed in this zone (Nitrosomonadaceae, Nitrospiraceae, and Nitrosococcaceae). The low abundance of methanotrophic microbes in the coastal sinkhole (mainly the cavern and cave zones) could be derived from the high concentrations of hydrogen sulfide previously reported at this location, which have been suggested to be toxic to methane-oxidizing microbes34. Therefore, a decrease in the anaerobic oxidation of methane, and a poor methane removal capacity is hypothesized in the sediments from this coastal sinkhole. Further research could focus on the influence of the saline intrusion on methanotrophic microbes and methane levels in El Zapote sinkhole sediments. As expected, photosynthetic bacterial abundances differed with the type of sinkhole and sediment zone. Both sinkholes are so the presence of daylight can start the photosynthetic process which would occur most in the WM zone. However, only the inland sinkhole showed a high abundance of photosynthetic bacteria within its WM zone. The coastal sinkhole water column and sediments would be deprived of photosynthetic bacteria since the water source is the underground aquifer, lacks photosynthesizers. Ammonia oxidizing bacteria (AOB) and archaea (AOA), and nitrite-oxidizing bacteria (NOB) were relatively abundant in the three sediment zones from the inland sinkhole and in the WM from the coastal sinkhole, these observation at the WM from El Zapote agree with previous observations2. Anaerobic ammonium oxidation (anammox) uses nitrite (as a product of nitrate reduction), as electron acceptor35. The high levels of nitrate concentration in the water column from the three sediment zones at the inland sinkhole and at the WM from the coastal sinkhole may influence the abundance of AOB, AOA and NOB in the sediments from these zones, while NH4+ and nitrite values were below detection limit ( More

1.Clobert, J., Danchin, E., Dhondt, A. A. & Nichols, J. D. Dispersal (Oxford University Press, 2001).
Google Scholar 
2.Ronce, O. How does it feel to be like a rolling stone? Ten questions about dispersal evolution. Annu. Rev. Ecol. Evol. Syst. 38, 231–253 (2007).
Google Scholar 
3.Clobert, J., Baguette, M., Benton, T. G. & Bullock, J. M. Dispersal Ecology and Evolution (Oxford University Press, 2012).
Google Scholar 
4.Cote, J., Fogarty, S., Brodin, T., Weinersmith, K. & Sih, A. Personality-dependent dispersal in the invasive mosquitofish: Group composition matters. Proc. R. Soc. B Biol. Sci. 278, 1670–1678 (2011).
Google Scholar 
5.Quinn, J. L., Cole, E. F., Patrick, S. C. & Sheldon, B. C. Scale and state dependence of the relationship between personality and dispersal in a great tit population. J. Anim. Ecol. 80, 918–928 (2011).PubMed 

Google Scholar 
6.Brodin, T., Lind, M. I., Wiberg, M. K. & Johansson, F. Personality trait differences between mainland and island populations in the common frog (Rana temporaria). Behav. Ecol. Sociobiol. 67, 135–143 (2013).
Google Scholar 
7.Wilson, D. S. Adaptive individual differences within single populations. Philos. Trans. R. Soc. London. Ser. B Biol. Sci. 353, 199–205 (1998).
Google Scholar 
8.Sih, A., Bell, A. & Johnson, J. C. Behavioral syndromes: An ecological and evolutionary overview. Trends Ecol. Evol. 19, 372–378 (2004).PubMed 

Google Scholar 
9.Sih, A., Bell, A. M., Johnson, J. C. & Ziemba, R. E. Behavioral syndromes: An integrative overview. Q. Rev. Biol. 79, 241–277 (2004).PubMed 

Google Scholar 
10.Réale, D., Reader, S. M., Sol, D., McDougall, P. T. & Dingemanse, N. J. Integrating animal temperament within ecology and evolution. Biol. Rev. 82, 291–318 (2007).PubMed 

Google Scholar 
11.Wolf, M. & Weissing, F. J. Animal personalities: Consequences for ecology and evolution. Trends Ecol. Evol. 27, 452–461 (2012).PubMed 

Google Scholar 
12.Juette, T., Cucherousset, J. & Cote, J. Animal personality and the ecological impacts of freshwater non-native species. Curr. Zool. 60, 417–427 (2014).
Google Scholar 
13.Duckworth, R. A. & Badyaev, A. V. Coupling of dispersal and aggression facilitates the rapid range expansion of a passerine bird. Proc. Natl. Acad. Sci. 104, 15017–15022 (2007).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
14.Conrad, J. L., Weinersmith, K. L., Brodin, T., Saltz, J. B. & Sih, A. Behavioural syndromes in fishes: A review with implications for ecology and fisheries management. J. Fish Biol. 78, 395–435 (2011).CAS 
PubMed 

Google Scholar 
15.Cote, J., Fogarty, S., Weinersmith, K., Brodin, T. & Sih, A. Personality traits and dispersal tendency in the invasive mosquitofish (Gambusia affinis). Proc. R. Soc. B Biol. Sci. 277, 1571–1579 (2010).
Google Scholar 
16.Malange, J., Izar, P. & Japyassú, H. Personality and behavioural syndrome in Necromys lasiurus (Rodentia: Cricetidae): Notes on dispersal and invasion processes. Acta Ethol. 19, 189–195 (2016).
Google Scholar 
17.Rees, E. M. A. et al. Socio-economic drivers of specialist anglers targeting the non-native European catfish (Silurus glanis) in the UK. PLoS ONE 12, e0178805 (2017).PubMed 
PubMed Central 

Google Scholar 
18.Bowler, D. E. & Benton, T. G. Causes and consequences of animal dispersal strategies: Relating individual behaviour to spatial dynamics. Biol. Rev. 80, 205–225 (2005).PubMed 

Google Scholar 
19.Clobert, J., Le Galliard, J.-F., Cote, J., Meylan, S. & Massot, M. Informed dispersal, heterogeneity in animal dispersal syndromes and the dynamics of spatially structured populations. Ecol. Lett. 12, 197–209 (2009).PubMed 

Google Scholar 
20.Dukes, J. S. & Mooney, H. A. Does global change increase the success of biological invaders?. Trends Ecol. Evol. 14, 135–139 (1999).CAS 
PubMed 

Google Scholar 
21.Gozlan, R. E., Britton, J. R., Cowx, I. & Copp, G. H. Current knowledge on non-native freshwater fish introductions. J. Fish Biol. 76, 751–786 (2010).
Google Scholar 
22.Pimentel, D. et al. Economic and environmental threats of alien plant, animal, and microbe invasions. Agric. Ecosyst. Environ. 84, 1–20 (2001).
Google Scholar 
23.Dingemanse, N. J., Kazem, A. J. N., Réale, D. & Wright, J. Behavioural reaction norms: Animal personality meets individual plasticity. Trends Ecol. Evol. 25, 81–89 (2010).PubMed 

Google Scholar 
24.Dochtermann, N. A., Schwab, T. & Sih, A. The contribution of additive genetic variation to personality variation: Heritability of personality. Proc. R. Soc. B Biol. Sci. 282, 20142201 (2015).
Google Scholar 
25.Duckworth, R. A. Evolution of personality: Developmental constraints on behavioral flexibility. Auk 127, 752–758 (2010).
Google Scholar 
26.Trillmich, F., Müller, T. & Müller, C. Understanding the evolution of personality requires the study of mechanisms behind the development and life history of personality traits. Biol. Lett. 14, 20170740 (2018).PubMed 
PubMed Central 

Google Scholar 
27.Dingemanse, N. J. & Réale, D. Natural selection and animal personality. Behaviour 142, 1159–1184 (2005).
Google Scholar 
28.Sih, A., Cote, J., Evans, M., Fogarty, S. & Pruitt, J. Ecological implications of behavioural syndromes. Ecol. Lett. 15, 278–289 (2012).PubMed 

Google Scholar 
29.Stamps, J. A. Growth-mortality tradeoffs and ‘personality traits’ in animals. Ecol. Lett. 10, 355–363 (2007).PubMed 

Google Scholar 
30.Chapple, D. G., Simmonds, S. M. & Wong, B. B. M. Can behavioral and personality traits influence the success of unintentional species introductions?. Trends Ecol. Evol. 27, 57–64 (2012).PubMed 

Google Scholar 
31.Hirsch, P. E., Thorlacius, M., Brodin, T. & Burkhardt-Holm, P. An approach to incorporate individual personality in modeling fish dispersal across in-stream barriers. Ecol. Evol. 7, 720–732 (2017).PubMed 

Google Scholar 
32.Groen, M. et al. Is there a role for aggression in round goby invasion fronts?. Behaviour 149, 685–703 (2012).
Google Scholar 
33.Urban, M. C., Phillips, B. L., Skelly, D. K. & Shine, R. A toad more traveled: The heterogeneous invasion dynamics of cane toads in Australia. Am. Nat. 171, E134–E148 (2008).PubMed 

Google Scholar 
34.Lopez, D. P., Jungman, A. A. & Rehage, J. S. Nonnative African jewelfish are more fit but not bolder at the invasion front: A trait comparison across an Everglades range expansion. Biol. Invasions 14, 2159–2174 (2012).
Google Scholar 
35.Dingemanse, N. J. & Wolf, M. Recent models for adaptive personality differences: A review. Philos. Trans. R. Soc. B Biol. Sci. 365, 3947–3958 (2010).
Google Scholar 
36.Dingemanse, N. J. & Réale, D. What is the evidence that natural selection maintains variation in animal personalities? In Animal Personalities: Behavior, Physiology, and Evolution (eds Carere, C. & Maestripieri, D.) 201–220 (University of Chicago Press, 2013).
Google Scholar 
37.Weiss, A. Personality traits: A view from the animal kingdom. J. Pers. 86, 12–22 (2018).PubMed 

Google Scholar 
38.Archard, G. A. & Braithwaite, V. A. The importance of wild populations in studies of animal temperament. J. Zool. 281, 149–160 (2010).
Google Scholar 
39.Holt, R. D., Keitt, T. H., Lewis, M. A., Maurer, B. A. & Taper, M. L. Theoretical models of species’ borders: Single species approaches. Oikos 108, 18–27 (2005).
Google Scholar 
40.Liedvogel, M., Chapman, B. B., Muheim, R. & Åkesson, S. The behavioural ecology of animal movement: Reflections upon potential synergies. Anim. Migr. 1, 39–46 (2013).
Google Scholar 
41.Campos-Candela, A., Palmer, M., Balle, S., Álvarez, A. & Alós, J. A mechanistic theory of personality-dependent movement behaviour based on dynamic energy budgets. Ecol. Lett. 22, 213–232 (2019).PubMed 

Google Scholar 
42.Bubb, D. H., Thom, T. J. & Lucas, M. C. Movement, dispersal and refuge use of co-occurring introduced and native crayfish. Freshw. Biol. 51, 1359–1368 (2006).
Google Scholar 
43.Luque, G. M. et al. The 100th of the world’s worst invasive alien species. Biol. Invasions 16, 981–985 (2014).
Google Scholar 
44.Galib, S. M., Findlay, J. S. & Lucas, M. C. Strong impacts of signal crayfish invasion on upland stream fish and invertebrate communities. Freshw. Biol. 66, 223–240 (2021).
Google Scholar 
45.Lindstrom, T., Brown, G. P., Sisson, S. A., Phillips, B. L. & Shine, R. Rapid shifts in dispersal behavior on an expanding range edge. Proc. Natl. Acad. Sci. 110, 13452–13456 (2013).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
46.Bubb, D. H., Thom, T. J. & Lucas, M. C. The within-catchment invasion of the non-indigenous signal crayfish Pacifastacus leniusculus (Dana), in upland rivers. Bull. Fr. Pêche Piscic. 376–377, 665–673 (2005).
Google Scholar 
47.Závorka, L., Lassus, R., Britton, J. R. & Cucherousset, J. Phenotypic responses of invasive species to removals affect ecosystem functioning and restoration. Glob. Chang. Biol. https://doi.org/10.1111/gcb.15271 (2020).Article 
PubMed 

Google Scholar 
48.Sbragaglia, V. & Breithaupt, T. Daily activity rhythms, chronotypes, and risk-taking behavior in the signal crayfish. Curr. Zool. https://doi.org/10.1093/cz/zoab023 (2021).Article 

Google Scholar 
49.R Core Team. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.r-project.org/ (2020).50.Pintor, L. M., Sih, A. & Bauer, M. L. Differences in aggression, activity and boldness between native and introduced populations of an invasive crayfish. Oikos 117, 1629–1636 (2008).
Google Scholar 
51.Rupia, E. J., Binning, S. A., Roche, D. G. & Lu, W. Fight-flight or freeze-hide? Personality and metabolic phenotype mediate physiological defence responses in flatfish. J. Anim. Ecol. 85, 927–937 (2016).PubMed 

Google Scholar 
52.Karavanich, C. & Atema, J. Individual recognition and memory in lobster dominance. Anim. Behav. 56, 1553–1560 (1998).CAS 
PubMed 

Google Scholar 
53.Houlihan, D., Govind, C. & El Haj, A. Energetics of swimming in Callinectes sapidus and walking in Homarus americanus. Comp. Biochem. Physiol. Part A Physiol. 82, 267–279 (1985).
Google Scholar 
54.Vogt, G. Functional anatomy. In Biology of Freshwater Crayfish (ed. Holdich, D. M.) 53–151 (Blackwell Science Ltd., 2002).
Google Scholar 
55.Southwood, T. R. E. & Henderson, P. A. Ecological Methods (Blackwell Science Ltd., 2000).
Google Scholar 
56.Clark, J. & Kershner, M. Size-dependent effects of visible implant elastomer marking on crayfish (Orconectes obscurus) growth, mortality, and tag retention. Crustaceana 79, 275–284 (2006).
Google Scholar 
57.Streissl, F. & Hödl, W. Habitat and shelter requirements of the stone crayfish, Austropotamobius torrentium Schrank. Hydrobiologia 477, 195–199 (2002).
Google Scholar 
58.Chadwick, D. D. A. et al. A novel ‘triple drawdown’ method highlights deficiencies in invasive alien crayfish survey and control techniques. J. Appl. Ecol. 58, 316–326 (2021).
Google Scholar 
59.Stoffel, M. A., Nakagawa, S. & Schielzeth, H. rptR: repeatability estimation and variance decomposition by generalized linear mixed-effects models. Methods Ecol. Evol. 8, 1639–1644 (2017).
Google Scholar 
60.Holm, S. A simple sequentially rejective multiple test procedure. Scand. J. Stat. 6, 65–70 (1979).MathSciNet 
MATH 

Google Scholar 
61.Quinn, G. P. & Keough, M. J. Experimental Design and Data Analysis for Biologists (Cambridge University Press, 2002).
Google Scholar 
62.Jackson, D. A. Stopping rules in principal components analysis: A comparison of heuristical and statistical approaches. Ecology 74, 2204–2214 (1993).
Google Scholar 
63.Budaev, S. V. Using principal components and factor analysis in animal behaviour research: Caveats and guidelines. Ethology 116, 472–480 (2010).
Google Scholar 
64.Robinson, C. A., Thom, T. J. & Lucas, M. C. Ranging behaviour of a large freshwater invertebrate, the white-clawed crayfish Austropotamobius pallipes. Freshw. Biol. 44, 509–521 (2000).
Google Scholar 
65.Bubb, D. H., O’Malley, O. J., Gooderham, A. C. & Lucas, M. C. Relative impacts of native and non-native crayfish on shelter use by an indigenous benthic fish. Aquat. Conserv. Mar. Freshw. Ecosyst. 19, 448–455 (2009).
Google Scholar 
66.Fox, J. & Weisberg, S. An R Companion to Applied Regression (Sage, 2011).
Google Scholar 
67.Burnham, K. P. & Anderson, D. R. Model Selection and Multimodel Inferences: A Practical Information-Theoretic Approach (Springer, 2002).MATH 

Google Scholar 
68.Bartoń, K. MuMIn: Multi-Model Inference. R Package version 1.43.6. (2019).69.Kleiber, C. & Zeileis, A. Applied Econometrics with R (Springer, 2008).MATH 

Google Scholar 
70.Edwards, D. D., Rapin, K. E. & Moore, P. A. Linking phenotypic correlations from a diverse set of laboratory tests to field behaviors in the crayfish, Orconectes virilis. Ethology 124, 311–330 (2018).
Google Scholar 
71.Teknomo, K. Similarity Measurements. https://people.revoledu.com/kardi/tutorial/Similarity (2015).72.Bell, A. M., Hankison, S. J. & Laskowski, K. L. The repeatability of behaviour: A meta-analysis. Anim. Behav. 77, 771–783 (2009).PubMed 
PubMed Central 

Google Scholar 
73.Vainikka, A., Rantala, M. J., Niemelä, P., Hirvonen, H. & Kortet, R. Boldness as a consistent personality trait in the noble crayfish, Astacus astacus. Acta Ethol. 14, 17–25 (2011).
Google Scholar 
74.Fraser, D. F., Gilliam, J. F., Daley, M. J., Le, A. N. & Skalski, G. T. Explaining leptokurtic movement distributions: Intrapopulation variation in boldness and exploration. Am. Nat. 158, 124–135 (2001).CAS 
PubMed 

Google Scholar 
75.Dingemanse, N. J., Both, C., van Noordwijk, A. J., Rutten, A. L. & Drent, P. J. Natal dispersal and personalities in great tits (Parus major). Proc. R. Soc. London. Ser. B Biol. Sci. 270, 741–747 (2003).
Google Scholar 
76.McMahon, T. E. & Tash, J. C. Experimental analysis of the role of emigration in population regulation of desert pupfish. Ecology 69, 1871–1883 (1988).
Google Scholar 
77.Porter, J. H. & Dooley, J. L. Animal dispersal patterns: A reassessment of simple mathematical models. Ecology 74, 2436–2443 (1993).
Google Scholar 
78.Einum, S., Sundt-Hansen, L. & Nislow, K. H. The partitioning of density-dependent dispersal, growth and survival throughout ontogeny in a highly fecund organism. Oikos 113, 489–496 (2006).
Google Scholar 
79.Lodge, D. M. & Hill, A. M. Factors governing species composition, population size and productivity of coolwater crayfishes. Nord. J. Freshw. Res. 69, 111–136 (1994).
Google Scholar 
80.Berthouly-Salazar, C., van Rensburg, B. J., Le Roux, J. J., van Vuuren, B. J. & Hui, C. Spatial sorting drives morphological variation in the invasive bird, Acridotheris tristis. PLoS ONE 7, e38145 (2012).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
81.Juanes, F. & Smith, L. D. The ecological consequences of limb damage and loss in decapod crustaceans: A review and prospectus. J. Exp. Mar. Biol. Ecol. 193, 197–223 (1995).
Google Scholar 
82.Wilshin, S. et al. Limping following limb loss increases locomotor stability. J. Exp. Biol. 221, jeb174268 (2018).PubMed 

Google Scholar 
83.Podgorniak, T., Blanchet, S., De Oliveira, E., Daverat, F. & Pierron, F. To boldly climb: Behavioural and cognitive differences in migrating European glass eels. R. Soc. Open Sci. 3, 150665 (2016).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
84.Bubb, D. H., Thom, T. J. & Lucas, M. C. Movement patterns of the invasive signal crayfish determined by PIT telemetry. Can. J. Zool. 84, 1202–1209 (2006).
Google Scholar 
85.Bilton, D. T., Freeland, J. R. & Okamura, B. Dispersal in freshwater invertebrates. Annu. Rev. Ecol. Syst. 32, 159–181 (2001).
Google Scholar 
86.Bubb, D. H., Thom, T. J. & Lucas, M. C. Movement and dispersal of the invasive signal crayfish Pacifastacus leniusculus in upland rivers. Freshw. Biol. 49, 357–368 (2004).
Google Scholar 
87.Hudina, S., Kutleša, P., Trgovčić, K. & Duplić, A. Dynamics of range expansion of the signal crayfish (Pacifastacus leniusculus) in a recently invaded region in Croatia. Aquat. Invasions 12, 67–75 (2017).
Google Scholar 
88.Wutz, S. & Geist, J. Sex- and size-specific migration patterns and habitat preferences of invasive signal crayfish (Pacifastacus leniusculus Dana). Limnologica 43, 59–66 (2013).
Google Scholar 
89.Fraser, H., Barnett, A., Parker, T. H. & Fidler, F. The role of replication studies in ecology. Ecol. Evol. 10, 5197–5207 (2020).PubMed 
PubMed Central 

Google Scholar 
90.Linzmaier, S. M., Goebel, L. S., Ruland, F. & Jeschke, J. M. Behavioral differences in an over-invasion scenario: marbled vs. spiny-cheek crayfish. Ecosphere 9, e02385 (2018).
Google Scholar 
91.Wang, X. et al. Anthropogenic habitat loss accelerates the range expansion of a global invader. Divers. Distrib. https://doi.org/10.1111/ddi.13359 (2021).Article 

Google Scholar  More

Relationships between microbial diversity and base flow indexThe number of reads obtained per sample and total number of OTUs obtained after rarefaction for each gene dataset are summarised in Table S2. According to taxonomic analyses of our 16S rRNA gene dataset, archaeal communities in our river sediment samples consisted largely of OTUs assigned to the Woesarchaeota (20.8% of OTUs and 24.7% of reads) and Methanomicrobia (16.9% of OTUs and 31.8% of reads). Of the functional groups analysed here, ten OTUs were assigned to AOA, Nitrososphaera (n = 8) and Nitrosopumilus (n = 2), that together formed 4.8% of all archaeal 16S rRNA reads. A total of 137 OTUs were assigned to orders of methanogenic archaea, with 15.3% and 16% of archaeal reads assigned to the orders Methanomicrobiales and Methanosarcinales, respectively, with other methanogen orders constituting a further 6.7% of reads.Bacterial communities were more diverse and OTUs assigned to taxa within the functional groups analysed here formed a relatively small proportion of our bacterial 16S rRNA gene dataset. Ammonia oxidising bacteria were represented by only five OTUs (all assigned to Nitrosospira) that together constituted 0.02% of the total bacterial community across our sediments. A further 84 OTUs were assigned to methanotrophic genera, and these OTUs contributed a total of 0.88% of all bacterial 16S rRNA sequences. These were Methylobacter (30 OTUs, 0.7% of bacterial sequences), Methylophilus (15 OTUs, 0.1% of bacterial sequences), Methylosoma (7 OTUs, 0.004% of bacterial sequences), Methylomonas and Methylotenera (6 OTUs each, 0.02 and 0.002% of bacterial sequences, respectively), and Methylosarcina (5 OTUs, 0.002% of bacterial sequences), with a further eight genera represented by a total of 15 OTUs. As reported previously, no OTUs were assigned to known anammox genera, which were likely below the limit of detection in our study [8].The OTU richness of archaeal communities (based on 16S rRNA amplicons) was negatively, albeit weakly, related to BFI (coef = 0.52, z = −2.95, adj-D2 = 0.12, P  More