Ecology

Building from this background the NAM took on these issues as its first-ever grand challenge, as a critical issue of import and urgency for us all. In 2018, the NAM empaneled an international, independent and multidisciplinary commission to create a global roadmap for healthy longevity, complete with evidence-based, targeted and actionable recommendations to move societies forward from an almost-exclusive focus on ‘coping with aging populations’ toward enabling individuals and societies to age successfully, and to reap the economic and societal benefits of longevity. The commission offers a way forward for governments and societies by beginning with recommendations for the next five years, and how these solutions can be financially sustainable through the creation of a virtuous cycle.To support these goals, the commission was to “(1) comprehensively address the challenges and opportunities presented by global aging population; (2) catalyze breakthrough ideas and research that will extend the human healthspan; and (3) generate transformative and scalable innovations world wide”8. The resulting comprehensive report, which was delayed in good measure by the COVID-19 pandemic, was released in June 2022 (ref. 8). We report here a summary of the high-level vision, goals, findings and recommendations of this global roadmap.The evidence for opportunities of longevity and the costs of inactionWe are seeing longer lives with increasing years spent in ill health (that is, the decompression of morbidity)9. The implications of longevity without health are costly ones for the individual, their families and for society. By contrast, scientific evidence shows that the majority of chronic diseases are preventable, and that prevention works at every age and stage of life. Further, the subset of individuals who are the beneficiaries of cumulative health-promoting conditions across the life course are demonstrating healthy longevity, defined as “the state in which years in good health approach the biological lifespan, with physical, cognitive and social functioning, enabling well-being across populations”8. However, only a minority of people in any country have the benefit of the necessary investments that promote health, and disparities in access to these investments across the life course are a major cause of unhealthy longevity. The costs of inaction in the face of widening disparities include the high risk of young people aging with more ill health, and the attendant costs to them and society.Further, the commission reports that when people have health and function in older age, the considerable cognitive and socioemotional capabilities and expertise that accrue with aging, and the prosocial goals of older age, constitute human and social capital assets that are unprecedented in both nature and scale. Contrary to disproven myths, workforce participation not only brings these valuable capabilities (such that intergenerational teams in the workplace are more productive and innovative than single-age-group teams), but older people working is also associated with more jobs for younger individuals10. In the USA and EU, it has been shown that older adults contribute 7% of gross domestic product (GDP) through paid work and the economic value of volunteering and caregiving11, even before opportunities are specifically expanded for the increasing older population. Societies that recognize this potential and invest to create both healthy longevity and the societal organizations and policies through which older adults can contribute to societal good will develop the opportunity for all ages to thrive. The return on investment will be to create older ages with health, function, dignity, meaning, purpose and opportunities — for those who desire it — to work longer, care for others or contribute in ways that they value to their community and future generations.The definition, principles and vision of ‘Vision 2050’ for healthy longevityThe global roadmap builds on the WHO ‘Decade of Healthy Ageing’, the UN Sustainable Development Goals for 2030 and other reports. It sets out principles for achieving healthy longevity using data and meaningful metrics to track achievement of outcomes and guide decision making. The report offers a vision empowered by the evidence: that, by 2050, societies will value the capabilities and assets of older people; all people will have the opportunity to live long lives with health and function; barriers to full participation by older people in society will have been solved; and that older people, with such health, will have the opportunity to engage in meaningful and productive activities. In turn, this societal engagement will create unprecedented social, human and economic capital, contributing to intergenerational well-being and cohesion, and to GDP.Implementing Vision 2050Accomplishing this vision demands ‘all-of-society’ intent — with aligned goals for healthy longevity and transformative action across public, private and academic sectors, and all of civil society and communities — and the implementation of evidence across the full and extending life course. Transforming only one component or sector (for example, health systems) will not be sufficient to create healthy longevity or its full opportunities. Rather, given that nations are complex systems, this vision for our future requires governmental leadership and transformation of all sectors of our complex societal system (Fig. 1).Fig. 1: Relevant actors for an all-of-society approach to healthy longevity.Healthy longevity requires government leadership and cooperation across all sectors. Adapted with permission from figure S-2 of ref. 8.Full size imageInvestment for healthy longevity — across the enabling sectors of health systems, social infrastructure and protections, the physical environment, and work and volunteering contributions — will require intentional planning and leadership to transform those components in tandem, and to resolve disrupters such as ageism, the social determinants of health and inequity, and pollution. These investments across all sectors will create the conditions for achieving healthy longevity and build new capital (human, social and economic) that will benefit all of society. As a result of these investments, society will see younger people thrive and move into a position to age with healthy longevity; those individuals who are already older will be recognized as valuable contributors to society in a ‘pay-it-forward’ stage of life. The underpinning social compact between citizens and government will support valuing each age group’s capabilities and goals, and the building of a society of well-being and cohesion across generations. This is at the center of the virtuous cycle for healthy longevity (Fig. 2)Fig. 2: The virtuous cycle of healthy longevity.Healthy longevity (top) is an outcome of a virtuous cycle, itself contributing to capital development (bottom left). Bottom right, capital (human, financial and social) supports enablers (work, physical environment, health systems and social infrastructure). The enablers propel the cycle, contributing to healthy longevity. Intentional investment for healthy longevity across all enabling sectors will create new capital that will benefit all of society. Adapted with permission from figure 1-4 of ref. 8.Full size imageGoals for initiating the transformation to healthy longevityThe commission identified the following changes that should occur from now to 2027 to start transformation of all of society, towards Vision 2050 and the creation of healthy longevity for all:

Creating social cohesion, social engagement and addressing the social determinants of health through social infrastructure are among the most effective determinants of slowed aging and the prevention of chronic conditions across the life course. Financial security in older age is essential for all.

Governments, the private sector and civil society should partner to design physical environments and infrastructure that are user-centered, and function as cohesion-enabling intergenerational communities for healthy longevity. Initiatives should focus on the inclusion of older people in the design, creating public spaces that promote social cohesion and intergenerational connection as well as mobility, physical activity and access to food, transportation, social services and engagement.

By 2027, governments should develop strategies and plans to arrive at adequately sized, geriatrically knowledgeable public health, clinical and long-term care workforces, and an integration of the pillars of the health system and social services. Together, these dimensions would foster and extend years of good health and support the diverse health needs and well-being of older people.

Governments should work to build the dividend of health longevity in collaboration with the business sector and civil society, to develop policies, incentives, and supportive systems that enable and encourage lifelong learning, and greater opportunities and necessary skills to engage in meaningful work or community volunteering across the lifespan.

We summarize the commission’s recommended goals for each of these sectors in brief in Box 1. Across all sectors, the key first steps that the commission identified are ones that can resolve obstacles to change and plan the change needed to shift multiple complex systems through both top-down and bottom-up approaches, in ways appropriate to each country and context. These initiatives should create enough momentum to foster early returns on investment and optimism to propel sustained investment for subsequent stages. This would need to begin for all governments by 2023, establishing calls to action to develop and implement data-driven, all-of-society plans to build the systems, policies, organizations and infrastructure needed, and for tracking change.Box 1 Goals for 2022–2027 to initiate the transformation to healthy longevityThese goals are reproduced from Global Roadmap for Healthy Longevity8.
Social infrastructure

Develop evidence-based multipronged strategies to reduce ageism against all groups.

Develop plans for ensuring basic financial security for all older people.

Develop strategies to increase financial literacy and mechanisms for promoting working longer, pension options and savings over the life course.

Plan opportunities for purposeful and meaningful engagement by older people at the family, community and societal levels.

Physical environment

At the societal level, improve broadband accessibility to reduce the digital divide and develop public transportation solutions that address first- and last-mile transportation.

At the city level, implement mitigation strategies to reduce the negative effects of the physical environment and related emergencies on older people (for example, air pollution and climate-induced events, including extreme heat and flooding) and design environments for connection and cohesion.

At the neighborhood level, promote and measure innovative policy solutions for healthy longevity, including affordable housing and intergenerational living, zoning and design for connection and cohesion, and the enabling of social capital.

At the home level, update physical infrastructure and policies to address affordability, provide coliving arrangements that match people’s goals and needs, and resolve insufficiencies and inefficiencies in housing stock.

Health systems

Establish healthy longevity as a major goal.

Increase investments in public health systems, which are needed to promote health and prevent disease, disability and injury at the population level, across the full life course. This may require rebalancing investments between this type of public health and medical care, recognizing that such public health is a public good and, as such, tends to be underinvested in.

Provide adequate primary care that includes preventive screening, addresses risk factors for chronic conditions and promotes positive health behaviors, and offers a continuum of medical care, including geriatrically knowledgeable care for older adults.

Make culturally sensitive, person-centered and equitable long-term care systems available, which (to the degree possible) offer dignity and honor people’s preferences about care settings.

Building the healthy longevity dividend

Governments, in collaboration with the business sector and civil society, should design (1) work environments and develop new policies that enable and encourage older adults who want or need to remain in the work force longer, and (2) engagement opportunities that strengthen communities at every stage of life.

Governments, employers and educational institutions should prioritize redesigning education systems to support lifelong learning and training, and invest in the science of learning and training for middle-aged and older adults.

Pilot innovations that incentivize and allow middle-aged and older adults to retool for multiple careers and/or participate as volunteers across their lifespan in roles with meaning and purpose. More

Arneth, A. et al. Terrestrial biogeochemical feedbacks in the climate system. Nat. Geosci. 3, 525–532 (2010).Article 

Google Scholar 
Adoption of the Paris Agreement FCCC/CP/2015/L.9/Rev.1 (UNFCC, 2015).IPCC Special Report on Climate Change and Land (eds Shukla, P. R. et al.) (IPCC, 2019).Oertel, C., Matschullat, J., Zurba, K., Zimmermann, F. & Erasmi, S. Greenhouse gas emissions from soils—a review. Geochemistry 76, 327–352 (2016).Article 

Google Scholar 
Schlesinger, W. H. & Bernhardt, E. S. Biogeochemistry: An Analysis of Global Change 3rd edn (Elsevier, 2013).Harris, N. L. et al. Global maps of twenty-first century forest carbon fluxes. Nat. Clim. Change 11, 234–240 (2021).Article 

Google Scholar 
Ackerman, D., Millet, D. B. & Chen, X. Global estimates of inorganic nitrogen deposition across four decades. Glob. Biogeochem. Cycles 33, 100–107 (2019).Article 

Google Scholar 
Du, E. Rise and fall of nitrogen deposition in the United States. Proc. Natl Acad. Sci. USA 113, E3594–E3595 (2016).Article 

Google Scholar 
Schmitz, A. et al. Responses of forest ecosystems in Europe to decreasing nitrogen deposition. Environ. Pollut. 244, 980–994 (2019).Article 

Google Scholar 
Hietz, P. et al. Long-term change in the nitrogen cycle of tropical forests. Science 334, 664–666 (2011).Article 

Google Scholar 
Fang, Y. T., Gundersen, P., Mo, J. M. & Zhu, W. X. Input and output of dissolved organic and inorganic nitrogen in subtropical forests of South China under high air pollution. Biogeosciences 5, 339–352 (2008).Article 

Google Scholar 
Yu, G. et al. Stabilization of atmospheric nitrogen deposition in China over the past decade. Nat. Geosci. 12, 424–429 (2019).Article 

Google Scholar 
Liu, L. L. & Greaver, T. L. A global perspective on belowground carbon dynamics under nitrogen enrichment. Ecol. Lett. 13, 819–828 (2010).Article 

Google Scholar 
LeBauer, D. S. & Treseder, K. K. Nitrogen limitation of net primary productivity in terrestrial ecosystems is globally distributed. Ecology 89, 371–379 (2008).Article 

Google Scholar 
Reich, P. B. et al. Scaling of respiration to nitrogen in leaves, stems and roots of higher land plants. Ecol. Lett. 11, 793–801 (2008).Article 

Google Scholar 
Cornwell, W. K. et al. Plant species traits are the predominant control on litter decomposition rates within biomes worldwide. Ecol. Lett. 11, 1065–1071 (2008).Article 

Google Scholar 
Mo, J. et al. Nitrogen addition reduces soil respiration in a mature tropical forest in southern China. Glob. Change Biol. 14, 403–412 (2008).Article 

Google Scholar 
Janssens, I. A. et al. Reduction of forest soil respiration in response to nitrogen deposition. Nat. Geosci. 3, 315–322 (2010).Article 

Google Scholar 
Zhong, Y., Yan, W. & Shangguan, Z. The effects of nitrogen enrichment on soil CO2 fluxes depending on temperature and soil properties. Glob. Ecol. Biogeogr. 25, 475–488 (2016).Article 

Google Scholar 
Deng, L. et al. Soil GHG fluxes are altered by N deposition: new data indicate lower N stimulation of the N2O flux and greater stimulation of the calculated C pools. Glob. Change Biol. 26, 2613–2629 (2020).Article 

Google Scholar 
Hagedorn, F., Kammer, A., Schmidt, M. W. I. & Goodale, C. L. Nitrogen addition alters mineralization dynamics of 13C-depleted leaf and twig litter and reduces leaching of older DOC from mineral soil. Glob. Change Biol. 18, 1412–1427 (2012).Article 

Google Scholar 
Du, Y. et al. Different types of nitrogen deposition show variable effects on the soil carbon cycle process of temperate forests. Glob. Change Biol. 20, 3222–3228 (2014).Article 

Google Scholar 
Yan, T. et al. Negative effect of nitrogen addition on soil respiration dependent on stand age: evidence from a 7-year field study of larch plantations in northern China. Agr. For. Meteorol. 262, 24–33 (2018).Article 

Google Scholar 
Xing, A. et al. Nonlinear responses of ecosystem carbon fluxes to nitrogen deposition in an old-growth boreal forest. Ecol. Lett. 25, 77–78 (2021).Article 

Google Scholar 
Melillo, J. et al. Long-term pattern and magnitude of soil carbon feedback to the climate system in a warming world. Science 358, 101–105 (2017).Article 

Google Scholar 
Gao, Q. et al. Stimulation of soil respiration by elevated CO2 is enhanced under nitrogen limitation in a decade-long grassland study. Proc. Natl Acad. Sci. USA 117, 33317–33324 (2020).Article 

Google Scholar 
Liu, X. J. et al. Nitrogen deposition and its ecological impact in China: an overview. Environ. Pollut. 159, 2251–2264 (2011).Article 

Google Scholar 
Zhu, F. F., Yoh, M., Gilliam, F. S., Lu, X. K. & Mo, J. M. Nutrient limitation in three lowland tropical forests in southern China receiving high nitrogen deposition: insights from fine root responses to nutrient additions. PLoS ONE 8, e82661 (2013).Article 

Google Scholar 
Wang, C. et al. Responses of soil microbial community to continuous experimental nitrogen additions for 13 years in a nitrogen-rich tropical forest. Soil Biol. Biochem. 121, 103–112 (2018).Article 

Google Scholar 
Priess, J. & Fölster, H. Microbial properties and soil respiration in submontane forests of Venezuelian Guyana: characteristics and response to fertilizer treatments. Soil Biol. Biochem. 33, 503–509 (2001).Article 

Google Scholar 
He, T., Wang, Q., Wang, S. & Zhang, F. Nitrogen addition altered the effect of belowground C allocation on soil respiration in a subtropical forest. PLoS ONE 11, e0155881 (2016).Article 

Google Scholar 
Fan, H. et al. Nitrogen deposition promotes ecosystem carbon accumulation by reducing soil carbon emission in a subtropical forest. Plant Soil 379, 361–371 (2014).Article 

Google Scholar 
Zheng, M. et al. Effects of nitrogen and phosphorus additions on nitrous oxide emission in a nitrogen-rich and two nitrogen-limited tropical forests. Biogeosciences 13, 3503–3517 (2016).Article 

Google Scholar 
Lu, X. et al. Nitrogen deposition accelerates soil carbon sequestration in tropical forests. Proc. Natl Acad. Sci. USA 118, e2020790118 (2021).Article 

Google Scholar 
Zhou, G. Y. et al. Old-growth forests can accumulate carbon in soils. Science 314, 1417–1417 (2006).Article 

Google Scholar 
Tian, J. et al. Long-term nitrogen addition modifies microbial composition and functions for slow carbon cycling and increased sequestration in tropical forest soil. Glob. Change Biol. 25, 3267–3281 (2019).Article 

Google Scholar 
Huang, N. et al. Spatial and temporal variations in global soil respiration and their relationships with climate and land cover. Sci. Adv. 6, eabb8508 (2020).Article 

Google Scholar 
Lu, X. K. et al. Effect of simulated N deposition on soil exchangeable cations in three forest types of subtropical China. Pedosphere 19, 189–198 (2009).Article 

Google Scholar 
Fang, Y., Gundersen, P., Mo, J. & Zhu, W. Nitrogen leaching in response to increased nitrogen inputs in subtropical monsoon forests in southern China. For. Ecol. Manage. 257, 332–342 (2009).Article 

Google Scholar 
Chen, X. M. et al. Effects of nitrogen deposition on soil organic carbon fractions in the subtropical forest ecosystems of S. China. J. Plant Nutr. Soil Sci. 175, 947–953 (2012).Article 

Google Scholar 
Fang, H. J. et al. 13C abundance, water-soluble and microbial biomass carbon as potential indicators of soil organic carbon dynamics in subtropical forests at different successional stages and subject to different nitrogen loads. Plant Soil 320, 243–254 (2009).Article 

Google Scholar 
Liu, L. et al. Effects of nitrogen and phosphorus additions on soil microbial biomass and community structure in two reforested tropical forests. Sci. Rep. 5, 14378–14378 (2014).Article 

Google Scholar 
Chen, H. et al. Nitrogen saturation in humid tropical forests after 6 years of nitrogen and phosphorus addition: hypothesis testing. Funct. Ecol. 30, 305–313 (2015).Article 

Google Scholar 
Lu, X., Mao, Q., Gilliam, F. S., Luo, Y. & Mo, J. Nitrogen deposition contributes to soil acidification in tropical ecosystems. Glob. Change Biol. 20, 3790–3801 (2014).Article 

Google Scholar 
Mao, Q. G. Impacts of Long-Term Nitrogen and Phosphorus Addition on Understory Plant Diversity in Subtropical Forests in Southern China. Doctoral Thesis, Univ. Chinese Academy of Sciences (2017).Xing, A. J. et al. High-level nitrogen additions accelerate soil respiration reduction over time in a boreal forest. Ecol. Lett. https://doi.org/10.1111/ele.14065 (2022).Cao, J. et al. Plant–bacteria–soil response to frequency of simulated nitrogen deposition has implications for global ecosystem change. Funct. Ecol. 34, 723–734 (2020).Article 

Google Scholar 
Mo, J. M., Brown, S., Peng, S. L. & Kong, G. H. Nitrogen availability in disturbed, rehabilitated and mature forests of tropical China. For. Ecol. Manage. 175, 573–583 (2003).Article 

Google Scholar 
Huang, Z. L., Ding, M. M., Zhang, Z. P. & Yi, W. M. The hydrological processes and nitrogen dynamics in a monsoon evergreen broad-leafed forest of Dinghushan. Acta Phytoecol. Sin. 18, 194–199 (1994).
Google Scholar 
Wright, R. F. & Rasmussen, L. Introduction to the NITREX and EXMAN projects. For. Ecol. Manage. 101, 1–7 (1998).Article 

Google Scholar 
Gundersen, P. et al. Impact of nitrogen deposition on nitrogen cycling in forests: a synthesis of NITREX data. For. Ecol. Manage. 101, 37–55 (1998).Article 

Google Scholar 
Aber, J. D. et al. Plant and soil responses to chronic nitrogen additions at the Harvard Forest, Massachusetts. Ecol. Appl. 3, 156–166 (1993).Article 

Google Scholar 
Cleveland, C. C. & Townsend, A. R. Nutrient additions to a tropical rain forest drive substantial soil carbon dioxide losses to the atmosphere. Proc. Natl Acad. Sci. USA 103, 10316–10321 (2006).Article 

Google Scholar 
Song, X. et al. Nitrogen addition increased CO2 uptake more than non-CO2 greenhouse gases emissions in a Moso bamboo forest. Sci. Adv. 6, eaaw5790 (2020).Article 

Google Scholar 
Lu, X. et al. Long-term nitrogen addition decreases carbon leaching in nitrogen-rich forest ecosystems. Biogeosciences 10, 3931–3941 (2013).Article 

Google Scholar 
Ackerman, D., Millet, D. B. & Chen, X. Global estimates of inorganic nitrogen deposition across four decades. Glob. Biogeochem. Cycles 33, 100–107 (2019).Article 

Google Scholar 
Tang, X., Liu, S., Zhou, G., Zhang, D. & Zhou, C. Soil–atmospheric exchange of CO2, CH4, and N2O in three subtropical forest ecosystems in southern China. Glob. Change Biol. 12, 546–560 (2006).Article 

Google Scholar 
Lei, J. et al. Temporal changes in global soil respiration since 1987. Nat. Commun. 12, 403 (2021).Article 

Google Scholar  More

Urban, M. C. Accelerating extinction risk from climate change. Science 348, 571–573 (2015).Article 
CAS 

Google Scholar 
Loarie, S. R. et al. The velocity of climate change. Nature 462, 1052–1055 (2009).Article 
CAS 

Google Scholar 
Pinsky, M. L., Eikeset, A. M., McCauley, D. J., Payne, J. L. & Sunday, J. M. Greater vulnerability to warming of marine versus terrestrial ectotherms. Nature 569, 108–111 (2019).Article 
CAS 

Google Scholar 
Hughes, A. R. et al. Predicting the sensitivity of marine populations to rising temperatures. Front. Ecol. Environ. 17, 17–24 (2019).Article 

Google Scholar 
Sunday, J. et al. Thermal tolerance patterns across latitude and elevation. Philos. Trans. R. Soc. B 374, 20190036 (2019).Article 

Google Scholar 
Bennett, S., Duarte, C. M., Marbà, N. & Wernberg, T. Integrating within-species variation in thermal physiology into climate change ecology. Philos. Trans. R. Soc. B 374, 20180550 (2019).Article 

Google Scholar 
Sasaki, M. C. & Dam, H. G. Integrating patterns of thermal tolerance and phenotypic plasticity with population genetics to improve understanding of vulnerability to warming in a widespread copepod. Glob. Change Biol. 25, 4147–4164 (2019).Article 

Google Scholar 
Kelly, M. W., Sanford, E. & Grosberg, R. K. Limited potential for adaptation to climate change in a broadly distributed marine crustacean. Proc. R. Soc. B 279, 349–356 (2012).Article 

Google Scholar 
Valladares, F. et al. The effects of phenotypic plasticity and local adaptation on forecasts of species range shifts under climate change. Ecol. Lett. 17, 1351–1364 (2014).Article 

Google Scholar 
Moran, E. V., Hartig, F. & Bell, D. M. Intraspecific trait variation across scales: implications for understanding global change responses. Glob. Change Biol. 22, 137–150 (2016).Article 

Google Scholar 
Razgour, O. et al. Considering adaptive genetic variation in climate change vulnerability assessment reduces species range loss projections. Proc. Natl Acad. Sci. USA 116, 10418–10423 (2019).Article 
CAS 

Google Scholar 
Seebacher, F., White, C. R. & Franklin, C. E. Physiological plasticity increases resilience of ectothermic animals to climate change. Nat. Clim. Change 5, 61–66 (2015).Article 

Google Scholar 
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).Article 
CAS 

Google Scholar 
Gunderson, A. R. & Stillman, J. H. Plasticity in thermal tolerance has limited potential to buffer ectotherms from global warming. Proc. R. Soc. B 282, 20150401 (2015).Article 

Google Scholar 
Barley, J. M. et al. Limited plasticity in thermally tolerant ectotherm populations: evidence for a trade-off. Proc. R. Soc. B 288, 202110765 (2021).Article 

Google Scholar 
Sunday, J. M., Bates, A. E. & Dulvy, N. K. Thermal tolerance and the global redistribution of animals. Nat. Clim. Change 2, 686–690 (2012).Article 

Google Scholar 
Grummer, J. A. et al. Aquatic landscape genomics and environmental effects on genetic variation. Trends Ecol. Evol. 34, 641–654 (2019).Article 

Google Scholar 
Kinlan, B. P. & Gaines, S. D. Propagule dispersal in marine and terrestrial environments: a community perspective. Ecology 84, 2007–2020 (2003).Article 

Google Scholar 
Lester, S. E., Ruttenberg, B. I., Gaines, S. D. & Kinlan, B. P. The relationship between dispersal ability and geographic range size. Ecol. Lett. 10, 745–758 (2007).Article 

Google Scholar 
Kinlan, B. P., Gaines, S. D. & Lester, S. E. Propagule dispersal and the scales of marine community process. Diversity Distrib. 11, 139–148 (2005).Article 

Google Scholar 
Mayr, E. Animal Species and Evolution (Harvard Univ. Press, 2014).Haldane, J. B. S. The relation between density regulation and natural selection. Proc. R. Soc. Lond. B 145, 306–308 (1956).Article 
CAS 

Google Scholar 
Marshall, D. J., Monro, K., Bode, M., Keough, M. J. & Swearer, S. Phenotype–environment mismatches reduce connectivity in the sea. Ecol. Lett. 13, 128–140 (2010).Article 
CAS 

Google Scholar 
Burgess, S. C., Treml, E. A. & Marshall, D. J. How do dispersal costs and habitat selection influence realized population connectivity? Ecology 93, 1378–1387 (2012).Article 

Google Scholar 
Sanford, E. & Kelly, M. W. Local adaptation in marine invertebrates. Annu. Rev. Mar. Sci. 3, 509–535 (2011).Article 

Google Scholar 
Caplat, P. et al. Looking beyond the mountain: dispersal barriers in a changing world. Front. Ecol. Environ. 14, 261–268 (2016).Article 

Google Scholar 
Nickols, K. J., Wilson White, J., Largier, J. L. & Gaylord, B. Marine population connectivity: reconciling large-scale dispersal and high self-retention. Am. Nat. 185, 196–211 (2015).Article 

Google Scholar 
Pinsky, M. L., Comte, L. & Sax, D. F. Unifying climate change biology across realms and taxa. Trends Ecol. Evol. https://doi.org/10.1016/j.tree.2022.04.011 (2022).Fourcade, Y. et al. Habitat amount and distribution modify community dynamics under climate change. Ecol. Lett. 24, 950–957 (2021).Article 

Google Scholar 
Kappes, H., Tackenberg, O. & Haase, P. Differences in dispersal- and colonization-related traits between taxa from the freshwater and the terrestrial realm. Aquat. Ecol. 48, 73–83 (2014).Article 
CAS 

Google Scholar 
Kinlan, B. P. & Gaines, S. D. Propagule dispersal in marine and terrestrial environments: a community perspective. Ecology 84, 2007–2020 (2003).Article 

Google Scholar 
Kappes, H. & Haase, P. Slow, but steady: dispersal of freshwater molluscs. Aquat. Sci. 74, 1–14 (2012).Article 

Google Scholar 
Sasaki, M. & Dam, H. G. Global patterns in copepod thermal tolerance. J. Plankton Res. 43, 598–609 (2021).Article 

Google Scholar 
Cereja, R. Critical thermal maxima in aquatic ectotherms. Ecol. Indic. 119, 106856 (2020).Article 

Google Scholar 
Vinagre, C. et al. Upper thermal limits and warming safety margins of coastal marine species – Indicator baseline for future reference. Ecol. Indic. 102, 644–649 (2019).Article 

Google Scholar 
Muñoz, M. M. The Bogert effect, a factor in evolution. Evolution 76, 49–66 (2022).Article 

Google Scholar 
Muñoz, M. M. & Bodensteiner, B. L. Janzen’s hypothesis meets the Bogert effect: connecting climate variation, thermoregulatory behavior, and rates of physiological evolution. Integr. Org. Biol. 1, oby002 (2019).Spence, A. R. & Tingley, M. W. The challenge of novel abiotic conditions for species undergoing climate-induced range shifts. Ecography 43, 1571–1590 (2020).Article 

Google Scholar 
Burrows, M. T. et al. The pace of shifting climate in marine and terrestrial ecosystems. Science 334, 652–655 (2011).Article 
CAS 

Google Scholar 
Steele, J. H., Brink, K. H. & Scott, B. E. Comparison of marine and terrestrial ecosystems: suggestions of an evolutionary perspective influenced by environmental variation. ICES J. Mar. Sci. 76, 50–59 (2019).Article 

Google Scholar 
Sexton, J. P., McIntyre, P. J., Angert, A. L. & Rice, K. J. Evolution and ecology of species range limits. Annu. Rev. Ecol. Evol. Syst. 40, 415–436 (2009).Article 

Google Scholar 
Chuang, A. & Peterson, C. R. Expanding population edges: theories, traits, and trade-offs. Glob. Change Biol. 22, 494–512 (2016).Article 

Google Scholar 
Bennett, J. M. et al. The evolution of critical thermal limits of life on Earth. Nat. Commun. 12, 1198 (2021).Article 
CAS 

Google Scholar 
Gaston, K. J. et al. Macrophysiology: a conceptual reunification. Am. Nat. 174, 595–612 (2009).Article 

Google Scholar 
Button, K. S. et al. Power failure: why small sample size undermines the reliability of neuroscience. Nat. Rev. Neurosci. 14, 365–376 (2013).Article 
CAS 

Google Scholar 
Gurevitch, J., Koricheva, J., Nakagawa, S. & Stewart, G. Meta-analysis and the science of research synthesis. Nature 555, 175–182 (2018).Article 
CAS 

Google Scholar 
Cooper, H., Hedges, L. V. & Valentine, J. C. The Handbook of Research Synthesis and Meta-Analysis (Russel Sage Foundation, 2009).Gleser, L. & Olkin, I. in The Handbook of Research Synthesis and Meta-Analysis (eds Cooper, H. et al.) Ch. 19 (Russel Sage Foundation, 2009).Huey, R. B., Hertz, P. E. & Sinervo, B. Behavioral drive versus behavioral inertia in evolution: a null model approach. Am. Nat. 161, 357–366 (2003).Article 

Google Scholar 
Bogert, C. M. Thermoregulation in reptiles, a factor in evolution. Evolution 3, 195–211 (1949).Article 
CAS 

Google Scholar 
Kearney, M., Shine, R. & Porter, W. P. The potential for behavioral thermoregulation to buffer ‘cold-blooded’’ animals against climate warming. Proc. Natl Acad. Sci. USA 10, 3835–3840 (2009).Article 

Google Scholar 
Denney, D. A., Jameel, M. I., Bemmels, J. B., Rochford, M. E. & Anderson, J. T. Small spaces, big impacts: contributions of micro-environmental variation to population persistence under climate change. AoB Plants 12, plaa005 (2020).Deutsch, C. A. et al. Impacts of climate warming on terrestrial ectotherms across latitude. Proc. Natl Acad. Sci. USA 105, 6668–6672 (2008).Article 
CAS 

Google Scholar 
Clusella-Trullas, S., Garcia, R. A., Terblanche, J. S. & Hoffmann, A. A. How useful are thermal vulnerability indices? Trends Ecol. Evol. 36, 1000–1010 (2021).Article 

Google Scholar 
Wanders, N., van Vliet, M. T. H., Wada, Y., Bierkens, M. F. P. & van Beek, L. P. H. High-resolution global water temperature modeling. Water Resour. Res. 55, 2760–2778 (2019).Article 

Google Scholar 
Todgham, A. E. & Stillman, J. H. Physiological responses to shifts in multiple environmental stressors: relevance in a changing world. Integr. Comp. Biol. 53, 539–544 (2013).Article 

Google Scholar 
Hoffmann, A. A. & Sgró, C. M. Climate change and evolutionary adaptation. Nature 470, 479–485 (2011).Article 
CAS 

Google Scholar 
Pespeni, M. H. & Palumbi, S. R. Signals of selection in outlier loci in a widely dispersing species across an environmental mosaic. Mol. Ecol. 22, 3580–3597 (2013).Article 
CAS 

Google Scholar 
Hoey, J. A. & Pinsky, M. L. Genomic signatures of environmental selection despite near-panmixia in summer flounder. Evolut. Appl. 11, 1732–1747 (2018).Article 
CAS 

Google Scholar 
Dirzo, R. et al. Defaunation in the Anthropocene. Science 345, 401–406 (2014).Article 
CAS 

Google Scholar 
Young, H. S., McCauley, D. J., Galetti, M. & Dirzo, R. Patterns, causes, and consequences of Anthropocene defaunation. Annu. Rev. Ecol. Evol. Syst. 47, 333–358 (2016).Article 

Google Scholar 
Morelli, T. L. et al. Managing Climate Change refugia for climate adaptation. PLoS ONE 11, e0159909 (2016).Article 

Google Scholar 
Cowen, R. K. & Sponaugle, S. Larval dispersal and marine population connectivity. Annu. Rev. Mar. Sci. 1, 443–466 (2009).Article 

Google Scholar 
Moher, D., Liberati, A., Tetzlaff, J., Altman, D. G. & PRISMA Group Preferred reporting items for systematic reviews and meta-analyses: the PRISMA statement. Ann. Internal Med. 151, 264–270 (2009).Article 

Google Scholar 
O’Dea, R. E. et al. Preferred reporting items for systematic reviews and meta-analyses in ecology and evolutionary biology: a PRISMA extension. Biol. Rev. https://doi.org/10.1111/brv.12721 (2021).Page, M. J. et al. The PRISMA 2020 statement: an updated guideline for reporting systematic reviews. BMJ 372, 89 (2021).Bennett, J. M. et al. GlobTherm, a global database on thermal tolerances for aquatic and terrestrial organisms. Sci. Data 5, 180022 1198 (2018).Lancaster, L. T. & Humphreys, A. M. Global variation in the thermal tolerances of plants. Proc. Natl Acad. Sci. USA 117, 13580–13587 (2020).Article 
CAS 

Google Scholar 
Rohatgi, A. WebPlotDigitizer (2020); https://automeris.io/WebPlotDigitizerAssis, J. et al. Bio-ORACLE v2.0: extending marine data layers for bioclimatic modelling. Glob. Ecol. Biogeogr. 27, 277–284 (2018).Article 

Google Scholar 
Karger, D. N. et al. Climatologies at high resolution for the Earth’s land surface areas. Sci. Data 4, 170122 (2017).Article 

Google Scholar 
Dee, D. P. et al. The ERA–interim reanalysis: configuration and performance of the data assimilation system. Q. J. R. Meteorolog. Soc. 137, 553–597 (2011).Article 

Google Scholar 
Helmuth, B. et al. Climate change and latitudinal patterns of intertidal thermal stress. Science 298, 1015–1017 (2002).Article 
CAS 

Google Scholar 
Helmuth, B. Thermal biology of rocky intertidal mussels: quantifying body temperature using climatological data. Ecology 80, 15–34 (1999).Article 

Google Scholar 
Bell, E. C. Environmental and morphological influences on thallus temperature and desiccation of the intertidal alga Mastocarpus papillatus Kützing. J. Exp. Mar. Biol. Ecol. 191, 29–55 (1995).Article 

Google Scholar 
R Core Team. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2020).Viechtbauer, W. Conducting meta-analyses in R with the metafor package. J. Stat. Software 36, 1–48 (2010).Article 

Google Scholar 
Sasaki, M. et al. Data for ‘greater local adaptation to temperature in the ocean than on land’. figshare https://doi.org/10.6084/m9.figshare.20173571 (2022). More

Research Group for Genomic Epidemiology, Technical University of Denmark, Kgs, Lyngby, DenmarkPatrick Munk, Christian Brinch, Frederik Duus Møller, Thomas N. Petersen, Rene S. Hendriksen, Anne Mette Seyfarth, Jette S. Kjeldgaard, Christina Aaby Svendsen & Frank M. AarestrupCentre for Immunity, Infection and Evolution, University of Edinburgh, Edinburgh, UKBram van Bunnik & Mark WoolhouseCentre for Antibiotic Resistance Research (CARe), University of Gothenburg, Gothenburg, SwedenFanny Berglund & D. G. Joakim LarssonDepartment of Viroscience, Erasmus MC, Rotterdam, The NetherlandsMarion KoopmansInstitute of Public Health, Tirana, AlbaniaArtan BegoUniversidad de Buenos Aires, Buenos Aires, ArgentinaPablo PowerMelbourne Water Corporation, Melbourne, AustraliaCatherine Rees & Kris CoventryCharles Darwin University, Darwin, AustraliaDionisia LambrinidisUniversity of Copenhagen, Frederiksberg C, DenmarkElizabeth Heather Jakobsen Neilson & Yaovi Mahuton Gildas HounmanouCharles Darwin University, Darwin Northern Territory, AustraliaKaren GibbCanberra Hospital, Canberra, AustraliaPeter CollignonALS Water, Scoresby, AustraliaSusan CassarAustrian Agency for Health and Food Safety (AGES), Vienna, AustriaFranz AllerbergerUniversity of Dhaka, Dhaka, BangladeshAnowara Begum & Zenat Zebin HossainEnvironmental Protection Department, Bridgetown, St. Michael, BarbadosCarlon WorrellLaboratoire Hospitalier Universitaire de Bruxelles (LHUB-ULB), Brussels, BelgiumOlivier VandenbergAQUAFIN NV, Aartselaar, BelgiumIlse PietersPolytechnic School of Abomey-Calavi, Abomey-Calavi, BeninDougnon Tamègnon VictorienUniversidad Catσlica Boliviana San Pablo, La Paz, BoliviaAngela Daniela Salazar Gutierrez & Freddy SoriaPublic Health Institute of the Republic of Srpska, Faculty of Medicine University of Banja Luka, Banja Luka, Bosnia and HerzegovinaVesna Rudić GrujićPublic Health Institute of the Republic of Srpska, Banja Luka, Bosnia and HerzegovinaNataša MazalicaBotswana International University of Science and Technology, Palapye, BotswanaTeddie O. RahubeUniversidade Federal de Minas Gerais, Belo Horizonte, BrazilCarlos Alberto Tagliati & Larissa Camila Ribeiro de SouzaOswaldo Cruz Institute, Rio de Janeiro, BrazilDalia RodriguesVale Institute of Technology, Belιm, PA, BrazilGuilherme OliveiraNational Center of Infectious and Parasitic Diseases, Sofia, BulgariaIvan IvanovUniversity of Ouagadougou, Ouagadougou, Burkina FasoBonkoungou Isidore Juste & Traoré OumarInstitut Pasteur du Cambodge, Phnom Penh, CambodiaThet Sopheak & Yith VuthyCentre Pasteur du Cameroun, Yaoundι, CameroonAntoinette Ngandijo, Ariane Nzouankeu & Ziem A. Abah Jacques OlivierUniversity of Regina, Regina, CanadaChristopher K. YostEau Terre Environnement Research Centre (INRS-ETE), Quebec City G1K 9A9, Canada and Indian Institute of Technology, Jammu, IndiaPratik KumarEau Terre Environnement Research Centre (INRS-ETE), Quebec City G1K 9A9, Canada and Lassonde School of Enginerring, York University, Toronto, CanadaSatinder Kaur BrarUniversity of N’Djamena, N’Djamena, ChadDjim-Adjim TaboEscuela de Medicina Veterinaria, Facultad de Ciencias de la Vida, Universidad Andres Bello, Santiago, ChileAiko D. AdellInstitute of Public Health, Santiago, ChileEsteban Paredes-Osses & Maria Cristina MartinezCentro de Biotecnologνa de los Recursos Naturales, Facultad de Ciencias Agrarias y Forestales, Talca, ChileSara Cuadros-OrellanaGuangdong Provincial Center for Disease Control and Prevention, Guangzhou, ChinaChangwen Ke, Huanying Zheng & Li BaishengThe Hong Kong Polytechnic University, Hong Kong, ChinaLok Ting Lau & Teresa ChungShantou University Medical College, Shantou, ChinaXiaoyang JiaoNanjing University of Information Science and Technology, Nanjing, ChinaYongjie YuCenter for Disease Control and Prevention of Henan province, Zhengzhou, ChinaZhao JiaYongColombian Integrated Program for Antimicrobial Resistance Surveillance – Coipars, CI Tibaitatα, Corporaciσn Colombiana de Investigaciσn Agropecuaria (AGROSAVIA), Tibaitatα – Mosquera, Cundinamarca, ColombiaJohan F. Bernal Morales, Maria Fernanda Valencia & Pilar Donado-GodoyInstitut Pasteur de Côte d’Ivoire, Abidjan, Côte d’IvoireKalpy Julien CoulibalyUniversity of Zagreb, Zagreb, CroatiaJasna HrenovicAndrija Stampar Teaching Institute of Public Health, Zagreb, CroatiaMatijana JergovićVeterinary Research Institute, Brno, Czech RepublicRenáta KarpíškováCentre de Recherche en Sciences Naturelles de Lwiro (CRSN-LWIRO), Bukavu, Democratic Republic of CongoZozo Nyarukweba DeogratiasBIOFOS A/S, Copenhagen K, DenmarkBodil ElsborgTechnical University of Denmark, Kgs., Lyngby, DenmarkLisbeth Truelstrup Hansen & Pernille Erland JensenSuez Canal University, Ismailia, EgyptMohamed AbouelnagaUniversity of Sadat City, Sadat City, EgyptMohamed Fathy SalemMinistry of Health, Environmental Microbiology, Tallinn, EstoniaMarliin KoolmeisterAddis Ababa University, Addis Ababa, EthiopiaMengistu Legesse & Tadesse EgualeUniversity of Helsinki, Helsinki, FinlandAnnamari HeikinheimoFrench Institute Search Pour L’exploitation De La Mer (Ifremer), Nantes, FranceSoizick Le Guyader & Julien SchaefferInstituto Nacional de Investigaciσn en Salud Pϊblica-INSPI (CRNRAM), Galαpagos, Quito, EcuadorJose Eduardo VillacisNational Public Health Laboratories, Ministry of Health and Social Welfare, Kotu, GambiaBakary SannehNational Center for Disease Control and Public Health, Tbilisi, GeorgiaLile MalaniaRobert Koch Institute, Berlin, GermanyAndreas Nitsche & Annika BrinkmannTechnische Universitδt Dresden, Institute of Hydrobiology, Dresden, GermanySara Schubert, Sina Hesse & Thomas U. BerendonkUniversity for Development Studies, Tamale, GhanaCourage Kosi Setsoafia SabaUniversity of Ghana, Accra, GhanaJibril MohammedKwame Nkrumah University of Science and Technology, Kumasi, PMB, GhanaPatrick Kwame FegloCouncil for Scientific and Industrial Research Water Research Institute, Accra, GhanaRegina Ama BanuVeterinary Research Institute of Thessaloniki, Hellenic Agricultural Organisation-DEMETER, Thermi, GreeceCharalampos KotzamanidisAthens Water Supply and Sewerage Company (EYDAP S.A.), Athens, GreeceEfthymios LytrasUniversidad de San Carlos de Guatemala, Guatemala City, GuatemalaSergio A. LickesSemmelweis University, Institute of Medical Microbiology, Budapest, HungaryBela KocsisUniversity of Veterinary Medicine, Budapest, HungaryNorbert SolymosiUniversity of Iceland, Reykjavνk, IcelandThorunn R. ThorsteinsdottirCochin University of Science and Technology, Cochin, IndiaAbdulla Mohamed HathaKasturba Medical College, Manipal, IndiaMamatha BallalApollo Diagnostics, Mangalore, IndiaSohan Rodney BangeraShiraz University of Medical Sciences, Shiraz, IranFereshteh FaniShahid Beheshti University of Medical Sciences, Tehran, IranMasoud AlebouyehNational University of Ireland Galway, Galway, IrelandDearbhaile Morris, Louise O’Connor & Martin CormicanBen Gurion University of the Negev and Ministry of Health, Beer-Sheva, IsraelJacob Moran-GiladIstituto Zooprofilattico Sperimentale del Lazio e della Toscana, Rome, ItalyAntonio Battisti, Elena Lavinia Diaconu & Patricia AlbaCNR – Water Research Institute, Verbania, ItalyGianluca Corno & Andrea Di CesareNational Institute of Infectious Diseases, Tokyo, JapanJunzo Hisatsune, Liansheng Yu, Makoto Kuroda, Motoyuki Sugai & Shizuo KayamaNational Center of Expertise, Taldykorgan, KazakhstanZeinegul ShakenovaMount Kenya University, Thika, KenyaCiira KiiyukiaKenya Medical Research Institute, Nairobi, KenyaEric Ng’enoUniversity of Prishtina “Hasan Prishtina” & National Institute of Public Health of Kosovo, Pristina, KosovoLul RakaKuwait Institute for Scientific Research, Kuwait City, KuwaitKazi Jamil, Saja Adel Fakhraldeen & Tareq AlaatiInstitute of Food Safety, Riga, LatviaAivars Bērziņš, Jeļena Avsejenko, Kristina Kokina, Madara Streikisa & Vadims BartkevicsAmerican University of Beirut, Beirut, LebanonGhassan M. MatarCentral Michigan University & Michigan Health Clinics, Saginaw, MI, USAZiad DaoudNational Food and Veterinary Risk Assessment Institute, Vilnius, LithuaniaAsta Pereckienė & Ceslova Butrimaite-AmbrozevicieneLuxembourg Institute of Science and Technology, Belvaux, LuxembourgChristian PennyInstitut Pasteur de Madagascar, Antananarivo, MadagascarAlexandra Bastaraud & Jean-Marc CollardUniversity of Antananarivo, Centre d’Infectiologie Charles Mιrieux, Antananarivo, MadagascarTiavina Rasolofoarison, Luc Hervé Samison & Mala Rakoto AndrianariveloUniversity of Malawi, Blantyre, MalawiDaniel Lawadi BandaMalaysian Genomics Resource Centre Berhad, Kuala Lumpur, MalaysiaArshana AminAIMST University, COMBio, Kedah, MalaysiaHeraa Rajandas & Sivachandran ParimannanWater Services Corporation, Luqa, MaltaDavid SpiteriEnvironmental Health Directorate, St. Venera, MaltaMalcolm Vella HaberUniversity of Mauritius, Reduit, MauritiusSunita J. SantchurnInstitute for Public Health Montenegro, Podgorica, MontenegroAleksandar Vujacic & Dijana DjurovicInstitut Pasteur du Maroc, Casablanca, MoroccoBrahim Bouchrif & Bouchra KarraouanCentro de Investigaηγo em Saϊde de Manhiηa (CISM), Maputo, MozambiqueDelfino Carlos VubilAgriculture and Forestry University, Kathmandu, NepalPushkar PalNational Institute for Public, Health and the Environment (RIVM), Bilthoven, The NetherlandsHeike Schmitt & Mark van PasselUniversity of Otago, Dunedin, New ZealandGert-Jan Jeunen & Neil GemmellUniversity of Otago, Christchurch, New ZealandStephen T. ChambersUniversity of Central America, Managua, NicaraguaFania Perez Mendoza & Jorge Huete-PιrezUniversidad Nacional Autσnoma de Nicaragua-Leσn, Leσn, NicaraguaSamuel VilchezUniversity of Ilorin, Ilorin, NigeriaAkeem Olayiwola Ahmed, Ibrahim Raufu Adisa & Ismail Ayoade OdetokunUniversity of Ibadan, Ibadan, NigeriaKayode FashaeNorwegian Institute of Public Health, Oslo, NorwayAnne-Marie Sørgaard & Astrid Louise WesterVEAS, Slemmestad, NorwayPia Ryrfors & Rune HolmstadUniversity of Agriculture, Faisalabad, PakistanMashkoor MohsinAga Khan University, Karachi, PakistanRumina Hasan & Sadia ShakoorLaboratorio Central de Salud Publica, Asuncion, ParaguayNatalie Weiler Gustafson & Claudia Huber SchillInstituto Nacional de Salud, Lima, PeruMaria Luz Zamudio RojasUniversidad de Piura, Piura, PeruJorge Echevarria Velasquez & Felipe Campos YauceWHO Environmental and Occupational Health, Manila, PhilippinesBonifacio B. MagtibayMaynilad Water Services, Inc., Quezon City, PhilippinesKris Catangcatang & Ruby SibuloNational Veterinary Research Institute, Pulawy, PolandDariusz WasylUniversidade Catσlica Portuguesa, CBQF – Centro de Biotecnologia e Quνmica Fina – Laboratσrio Associado, Escola Superior de Biotecnologia, Porto, PortugalCelia Manaia & Jaqueline RochaAguas do Tejo Atlantico, Lisboa, PortugalJose Martins & Pedro ÁlvaroGwangju Institute of Science and Technology, Gwangju, Republic of KoreaDoris Di Yoong Wen, Hanseob Shin & Hor-Gil HurKorea Advanced Institute of Science and Technology, Daejeon, Republic of KoreaSukhwan YoonInstitute of Public Health of the Republic of North Macedonia, Skopje, Republic of North MacedoniaGolubinka Bosevska & Mihail KochubovskiState Medical and Pharmaceutical University, Chișinău, Republic of MoldovaRadu CojocaruNational Agency for Public Health, Chișinău, Republic of MoldovaOlga BurduniucKing Abdullah University of Science and Technology, Thuwal, Saudi ArabiaPei-Ying HongUniversity of Edinburgh, Edinburgh, Scotland, UKMeghan Rose PerryInstitut Pasteur de Dakar, Dakar, SenegalAmy GassamaInstitute of Veterinary Medicine of Serbia, Belgrade, SerbiaVladimir RadosavljevicNanyang Technological University, Singapore, SingaporeMoon Y. F. Tay, Rogelio Zuniga-Montanez & Stefan WuertzPublic Health Authority of the Slovak Republic, Bratislava, SlovakiaDagmar Gavačová, Katarína Pastuchová & Peter TruskaNational Laboratory of Health, Environment and Food, Ljubljana, SloveniaMarija TrkovIndependent consultant, Johannesburg, South AfricaKaren KeddyDaspoort Waste Water Treatment Works, Pretoria, South AfricaKerneels EsterhuyseKorea Advanced Institute of Science and Technology, Daejeon, South KoreaMin Joon SongSchool of Veterinary Sciences, Lugo, SpainMarcos Quintela-BalujaLabaqua, Santiago de Compostela, SpainMariano Gomez LopezIRTA, Centre de Recerca en Sanitat Animal (CReSA, IRTA-UAB), Campus de la Universitat Autonoma de Barcelona, Bellaterra, SpainMarta Cerdà-CuéllarUniversity of Kelaniya, Ragama, Sri LankaR. R. D. P. Perera, N. K. B. K. R. G. W. Bandara & H. I. PremasiriMedical Research Institute, Colombo, Sri LankaSujatha PathirageCaribbean Public Health Agency, Catries, Saint LuciaKareem CharlemagneThe Sahlgrenska Academy at the University of Gothenburg, Gothenburg, SwedenCarolin RutgerssonSwedish University of Agricultural Sciences, Uppsala, SwedenLeif Norrgren & Stefan ÖrnFederal Food Safety and Veterinary Office (FSVO), Bern, SwitzerlandRenate BossAra Region Bern AG, Herrenschwanden, SwitzerlandTanja Van der HeijdenCenters for Disease Control, Taipei, TaiwanYu-Ping HongKilimanjaro Clinical Research Institute, Moshi, TanzaniaHappiness Houka KumburuSokoine University of Agriculture, Morogoro, TanzaniaRobinson Hammerthon MdegelaFaculty of Science and Technology, Suratthani Rajabhat University, Surat Thani, ThailandKaknokrat ChonsinFaculty of Public Health, Mahidol University, Bangkok, ThailandOrasa SuthienkulFaculty of Medicine Siriraj Hospital, Bangkok, ThailandVisanu ThamlikitkulNational Institute for Public Health and the Environment (RIVM), Bilthoven, NetherlandsAna Maria de Roda HusmanNational Institute of Hygiene, Lomι, TogoBawimodom BidjadaAgence de Mιdecine Prιventive, Dapaong, TogoBerthe-Marie Njanpop-LafourcadeDivision of Integrated Surveillance of Health Emergencies and Response, Lomι, TogoSomtinda Christelle Nikiema-PessinabaPublic Health Institution of Turkey, Ankara, TurkeyBelkis LeventHatay Mustafa Kemal University, Hatay, TurkeyCemil KurekciMakerere University, Kampala, UgandaFrancis Ejobi & John Bosco KaluleAbu Dhabi Public Health Center, Abu Dhai, United Arab EmiratesJens ThomsenDubai municipality, WWTP Al Aweer, Dubai, UAEOuidiane ObaidiRashid Hospital, Dubai, UAELaila Mohamed JassimNorthumbrian Water, Northumbria House, Abbey Road, Pity Me, Durham, UKAndrew MooreUniversity of Exeter Medical School, Cornwall, UKAnne Leonard, Lihong Zhang & William H. GazeNewcastle University, Newcastle upon Tyne, UKDavid W. Graham & Joshua T. BunceBrightwater Treatment Plant, Woodinville, WA, USABrett LeforDepartment of Environmental Sciences and Engineering, University of North Carolina at Chapel Hill, Chapel Hill, NC, USADrew Capone & Joe BrownUniversity of North Carolina, Chapel Hill, USAEmanuele Sozzi & Mark D. SobseyUniversity of Washington, Seattle, WA, USAJohn Scott Meschke, Nicola Koren Beck, Pardi Sukapanpatharam & Phuong TruongBaylor University, Waco, USAMichael DavisColumbia Boulevard WWTP, Portland, USARonald LilienthalEastern Illinois University, Charleston, USASanghoon KangThe Ohio State University, Columbus Ohio, USAThomas E. WittumLaboratorio Tecnolσgico del Uruguay, Montevideo, UruguayNatalia Rigamonti & Patricia BaklayanInstitute of Public Health in Ho Chi Minh City, Ho Chi Minh, VietnamChinh Dang Van, Doan Minh Nguyen Tran & Nguyen Do PhucUniversity of Zambia, Lusaka, ZambiaGeoffrey KwendaF.M.A., M.K., and M.W. conceived the study and secured funding. R.S.H., A.M.S., C.A.A.S., and J.S.K. organized sample collection, material transfer, and logistics. F.D.M., P.M., and C.B. did quality control, sample selection, and outlier detection. P.M., C.B., F.D.M., T.N.P., and F.B. performed bioinformatics analyses. P.M. and C.B. carried out data and statistical analyses and visualization. P.M. and F.M.A. drafted the initial manuscript with input from C.B., B.v.B., D.G.J.L., M.W., and M.K. The Global Sewage Consortium authors carried out sewage sampling, filled in metadata and shipped the samples to DTU. All authors helped to review and improve the manuscript. More

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.This is a summary of: Sasaki, M. et al. Greater evolutionary divergence of thermal limits within marine than terrestrial species. Nat. Clim. Change https://doi.org/10.1038/s41558-022-01534-y (2022). More

Montgomery, D. R. Soil erosion and agricultural sustainability. Proc. Natl. Acad. Sci. 104, 13268–13272 (2007).Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Evans, D. L., Quinton, J. N., Davies, J. A. C., Zhao, J. & Govers, G. Soil lifespans and how they can be extended by land use and management change. Environ. Res. Lett. 15, 1. https://doi.org/10.1088/1748-9326/aba2fd (2020).Adhikari, K. & Hartemink, A. E. Linking soils to ecosystem services—A global review. Geoderma 262, 101–111. https://doi.org/10.1016/j.geoderma.2015.08.009 (2016).Article 
CAS 

Google Scholar 
Gao, Y. et al. Effects of tillage methods on soil carbon and wind erosion. Land Degrad. Dev. 27, 583–591. https://doi.org/10.1002/ldr.2404 (2016).Article 

Google Scholar 
Klik, A. & Rosner, J. Long-term experience with conservation tillage practices in Austria: Impacts on soil erosion processes. Soil Till. Res. 203, 1. https://doi.org/10.1016/j.still.2020.104669 (2020).Seitz, S. et al. Conservation tillage and organic farming reduce soil erosion. Agron. Sustain. Dev. 39, 1. https://doi.org/10.1007/s13593-018-0545-z (2018).Lal, R., Reicosky, D. C. & Hanson, J. D. Evolution of the plow over 10,000 years and the rationale for no-till farming. Soil Till. Res. 93, 1–12. https://doi.org/10.1016/j.still.2006.11.004 (2007).Article 

Google Scholar 
Mal, P., Schmitz, M. & Hesse, J. W. Economic and environmental effects of conservation tillage with glyphosate use: A case study of Germany. Outlooks Pest Manag. 26, 24–27. https://doi.org/10.1564/v26_feb_07 (2015).Article 

Google Scholar 
Statistisches Bundesamt. Land- und Forstwirtschaft, Fischerei. Bodenbearbeitung, Bewässerung, Landschaftselemente. Erhebung über landwirtschaftliche Produktionsmethoden (ELPM). 2010. (2011).Quinton, J. N., Govers, G., Van Oost, K. & Bardgett, R. D. The impact of agricultural soil erosion on biogeochemical cycling. Nat. Geosci. 3, 311–314. https://doi.org/10.1038/ngeo838 (2010).Article 
CAS 

Google Scholar 
Öttl, L. K. et al. Tillage erosion as an important driver of in-field biomass patterns in an intensively used hummocky landscape. Land Degrad. Dev. 32, 3077–3091. https://doi.org/10.1002/ldr.3968 (2021).Article 

Google Scholar 
Wilken, F., Ketterer, M., Koszinski, S., Sommer, M. & Fiener, P. Understanding the role of water and tillage erosion from 239+240Pu tracer measurements using inverse modelling. SOIL 6, 549–564. https://doi.org/10.5194/soil-6-549-2020 (2020).Article 
CAS 

Google Scholar 
Van Oost, K., Govers, G., De Alba, S. & Quine, T. A. Tillage erosion: A review of controlling factors and implications for soil quality. Prog. Phys. Geogr. 30, 443–466. https://doi.org/10.1191/0309133306pp487ra (2006).Article 

Google Scholar 
Winnige, B. Ergebnisse zur Bodenverlagerung durch Bearbeitungserosion in der Jungmoränenlandschaft Nordostdeutschlands—Investigations of soil movement by tillage as a type of soil erosion in the young moraine soil landscape of Northeast Germany. Arch. Agron. Soil Sci. 50, 319–327. https://doi.org/10.1080/03650340410001663864 (2004).Article 

Google Scholar 
Fiener, P., Wilken, F. & Auerswald, K. Filling the gap between plot and landscape scale—eight years of soil erosion monitoring in 14 adjacent watersheds under soil conservation at Scheyern, Southern Germany. Adv. Geosci. 48, 31–48. https://doi.org/10.5194/adgeo-48-31-2019 (2019).Article 

Google Scholar 
Fiener, P. et al. Uncertainties in assessing tillage erosion—How appropriate are our measuring techniques?. Geomorphology 304, 214–225. https://doi.org/10.1016/j.geomorph.2017.12.031 (2018).Article 

Google Scholar 
Kimaro, D. N., Deckers, J. A., Poesen, J., Kilasara, M. & Msanya, B. M. Short and medium term assessment of tillage erosion in the Uluguru Mountains Tanzania. Soil Till. Res. 81, 97–108. https://doi.org/10.1016/j.still.2004.05.006 (2005).Article 

Google Scholar 
Sadowski, H. & Sorge, B. Der Normalhöhenpunkt von 1912 – Datumspunkt des DHHN 2012? Vermessung Brandenburg (2005).Lobb, D. A., Kachanoski, R. G. & Miller, M. H. Tillage translocation and tillage erosion in the complex upland landscapes of southwestern Ontario Canada. Soil Till. Res. 51, 1. https://doi.org/10.1016/S0167-1987(99)00037-9 (1999).Article 

Google Scholar 
Zhang, J. H. & Li, F. C. An appraisal of two tracer methods for estimating tillage erosion rates under hoeing tillage. Proc. Environ. Sci. 11, 1227–1233. https://doi.org/10.1016/j.proenv.2011.12.184 (2011).Article 

Google Scholar 
Turkelboom, F. et al. Assessment of tillage erosion rates on steep slopes in northern Thailand. CATENA 29, 29–44 (1997).Article 
CAS 

Google Scholar 
Van Muysen, W., Govers, G., Van Oost, K. & Van Rompaey, A. The effect of tillage depth, tillage speed, and soil condition on chisel tillage erosivity. J. Soil Water Conserv. 55, 355–364 (2000).
Google Scholar 
Quine, T. A., Desmet, P. J. J., Govers, G., Vandaele, K. & Walling, D. E. A comparison of the roles of tillage and water erosion in landform development and sediment export on agricultural land near Leuven, Belgium. IAHS Publ. 224, 77–86 (1994).CAS 

Google Scholar 
Heckrath, G. et al. Tillage erosion and its effect on soil properties and crop yield in Denmark. J. Environ. Qual. 34, 312–324. https://doi.org/10.2134/jeq2005.0312a (2005).Article 
CAS 
PubMed 

Google Scholar 
Carter, M. R. Conservation tillage. Encyclop. Soils Environ. 1, 306–311. https://doi.org/10.1016/B0-12-348530-4/00270-8 (2005).Article 

Google Scholar 
Govers, G., Vandaele, K., Desmet, P., Poesen, J. & Bunte, K. The role of tillage in soil redistribution on hillslopes. Eur. J. Soil Sci. 45, 469–478. https://doi.org/10.1111/j.1365-2389.1994.tb00532.x (1994).Article 

Google Scholar 
Marques da Silva, J. R. & Alexandre, C. Soil carbonation processes as evidence of tillage-induced erosion. Soil Till. Res. 78, 217–224. https://doi.org/10.1016/j.still.2004.02.008 (2004).Mech, S. J. & Free, G. R. Movement of soil during tillage operations. Agric. Eng. 1, 379–382 (1942).
Google Scholar 
Tiessen, K. H. D., Mehuys, G. R., Lobb, D. A. & Rees, H. W. Tillage erosion within potato production systems in Atlantic Canada: I. Measurement of tillage translocation by implements used in seedbed preparation. Soil Till. Res. 95, 308–319. https://doi.org/10.1016/j.still.2007.02.003 (2007).Article 

Google Scholar 
Marques da Silva, J. R., Soares, J. M. C. N. & Karlen, D. L. Implement and soil condition effects on tillage-induced erosion. Soil Till. Res. 78, 207–216. https://doi.org/10.1016/j.still.2004.02.009 (2004).Article 

Google Scholar 
Kietzer, B. Aufklärung der Bodenverlagerung durch Bearbeitungserosion in Jungmoränenlandschaften—Elucidation of soil displacement by tillage erosion in young moraine landscapes PhD thesis, Technical University of Berlin, (2007).Lüthgens, C., Böse, M. & Preusser, F. Age of the Pomeranian ice-marginal position in northeastern Germany determined by Optically Stimulated Luminescence (OSL) dating of glaciofluvial sediments. Boreas 40, 598–615. https://doi.org/10.1111/j.1502-3885.2011.00211.x (2011).Article 

Google Scholar 
Deumlich, D., Schmidt, R. & Sommer, M. A multiscale soil-landform relationship in the glacial-drift area based on digital terrain analysis and soil attributes. J. Plant Nutr. Soil Sci. 173, 843–851. https://doi.org/10.1002/jpln.200900094 (2010).Article 
CAS 

Google Scholar 
Koszinski, S., Gerke, H. H., Hierold, W. & Sommer, M. Geophysical-based modeling of a kettle hole catchment of the morainic soil landscape. Vadose Zone J. 12, 1. https://doi.org/10.2136/vzj2013.02.0044 (2013).Article 

Google Scholar 
Sommer, M., Gerke, H. H. & Deumlich, D. Modelling soil landscape genesis: A “time split” approach for hummocky agricultural landscapes. Geoderma 145, 480–493. https://doi.org/10.1016/j.geoderma.2008.01.012 (2008).Article 
CAS 

Google Scholar 
DWD Climate Data Center (CDC). Historical hourly station observations of 2m air temperature and humidity for Germany, version v006. (2018).DWD Climate Data Center (CDC). Historical hourly station observations of precipitation for Germany, version v21.3. (2021).Zhang, H. et al. Evaluating the potential of post-processing kinematic (PPK) georeferencing for UAV-based structure- from-motion (SfM) photogrammetry and surface change detection. Earth Surf. Dyn. 7, 807–827. https://doi.org/10.5194/esurf-7-807-2019 (2019).Article 

Google Scholar 
Lindstrom, M. J., Nelson, W. W., Schumacher, T. E. & Lemme, G. D. Soil movement by tillage as affected by slope. Soil Till. Res. 17, 255–264. https://doi.org/10.1016/0167-1987(90)90040-K (1990).Article 

Google Scholar 
Crawley, M. J. The R book. 2nd edn, (Wiley, 2013).Wickham, H. ggplot2: Elegant graphics for data analysis (Springer, 2016).Book 
MATH 

Google Scholar 
R Core Team. A language and environment for statistical computing. (2021).De Alba, S. Modelling the effects of complex topography and patterns of tillage on soil translocation by tillage with mouldboard plough. J. Soil Water Conserv. 1, 335–345 (2001).
Google Scholar 
Gerontidis, D. V. S. et al. The effect of moldboard plow on tillage erosion along a hillslope. J. Soil Water Conserv. 56, 147–152 (2001).
Google Scholar 
Heckrath, G., Halekoh, U., Djurhuus, J. & Govers, G. The effect of tillage direction on soil redistribution by mouldboard ploughing on complex slopes. Soil Tillage Res. 88, 225–241. https://doi.org/10.1016/j.still.2005.06.001 (2006).Article 

Google Scholar 
Kosmas, C. et al. The effects of tillage displaced soil on soil properties and wheat biomass. Soil Till Res. 58, 31–44. https://doi.org/10.1016/S0167-1987(00)00175-6 (2001).Article 

Google Scholar 
Lindstrom, M. J., Nelson, W. W. & Schumacher, T. E. Quantifying tillage erosion rates due to moldboard plowing. Soil Till Res. 24, 243–255. https://doi.org/10.1016/0167-1987(92)90090-X (1992).Article 

Google Scholar 
Lobb, D. A., Kachanoski, R. G. & Miller, M. H. Tillage translocation and tillage erosion on shoulder slope landscape positions measured using 137Cs as a tracer. Can. J. Soil Sci. 75, 211–218. https://doi.org/10.4141/cjss95-029 (1995).Article 

Google Scholar 
Quine, T. A. & Zhang, Y. Re-defining tillage erosion: Quantifying intensity–direction relationships for complex terrain: 1. Derivation of an adirectional soil transport coefficient. Soil Use Manag. 20, 114–123. https://doi.org/10.1111/j.1475-2743.2004.tb00346.x (2004).Article 

Google Scholar 
Quine, T. A., Basher, L. R. & Nicholas, A. P. Tillage erosion intensity in the South Canterbury Downlands, New Zealand. Aust. J. Soil Res. 41, 789–807. https://doi.org/10.1071/SR02063 (2003).Article 

Google Scholar 
Revel, J. C. & Guiresse, M. Erosion due to cultivation of calcareous clay soils on the hillsides of south west France: I. Effect of former farming practices. Soil Till Res. 35, 147–155. https://doi.org/10.1016/0167-1987(95)00482-3 (1995).Article 

Google Scholar 
Van Muysen, W. & Govers, G. Soil displacement and tillage erosion during secondary tillage operations: The case of rotary harrow and seeding equipment. Soil Till Res. 65, 185–191. https://doi.org/10.1016/S0167-1987(01)00284-7 (2002).Article 

Google Scholar 
Van Muysen, W., Govers, G., Bergkamp, G., Roxo, M. & Poesen, J. Measurement and modelling of the effects of initial soil conditions and slope gradient on soil translocation by tillage. Soil Till Res. 51, 303–316. https://doi.org/10.1016/S0167-1987(99)00044-6 (1999).Article 

Google Scholar 
Poesen, J. et al. Patterns of rock fragment cover generated by tillage erosion. Geomorphology 18, 183–197. https://doi.org/10.1016/S0169-555X(96)00025-6 (1997).Article 

Google Scholar 
Quine, T. A. et al. Fine-earth translocation by tillage in stony soils in the Guadalentin, south-east Spain: An investigation using caesium-134. Soil Till Res. 51, 279–301. https://doi.org/10.1016/S0167-1987(99)00043-4 (1999).Article 
MathSciNet 

Google Scholar 
Kemper, W. D. & Rosenau, R. C. Soil cohesion as affected by time and water content. Soil Sci. Soc. Am. J. 1, 1001–1006. https://doi.org/10.2136/sssaj1984.03615995004800050009x (1984).Article 

Google Scholar 
Reinermann, S., Gessner, U., Asam, S., Kuenzer, C. & Dech, S. The effect of droughts on vegetation condition in Germany: An analysis based on two decades of satellite earth observation time series and crop yield statistics. Rem. Sens. 11, 1. https://doi.org/10.3390/rs11151783 (2019).Article 

Google Scholar 
Lüttger, A. B. & Feike, T. Development of heat and drought related extreme weather events and their effect on winter wheat yields in Germany. Theor. Appl. Climatol. 1, 15–29. https://doi.org/10.1007/s00704-017-2076-y (2018).Article 

Google Scholar 
Madarász, B. et al. Conservation tillage vs. conventional tillage: Long-term effects on yields in continental, sub-humid Central Europe. Hungary. Int. J. Agric. Sustain. 14, 408–427. https://doi.org/10.1080/14735903.2016.1150022 (2016).Article 

Google Scholar 
Lowder, S. K., Skoet, J. & Raney, T. The number, size, and distribution of farms, smallholder farms, and family farms worldwide. World Dev. 87, 16–29. https://doi.org/10.1016/j.worlddev.2015.10.041 (2016).Article 

Google Scholar 
Napoli, M., Altobelli, F. & Orlandini, S. Effect of land set up systems on soil losses. Ital. J. Agron. 15, 306–314. https://doi.org/10.4081/ija.2020.1768 (2020).Article 

Google Scholar 
Dumanski, J., Peiretti, R., Benites, J. R., McGarry, D. & Pieri, C. The paradigm of conservation agriculture. In Proceedings of World Association of Soil and Water Conservation, 58–64 (2006). More

Ebenhard, T. Colonization in metapopulations: A review of theory and observations. Biol. J. Linn. Soc. 42, 105–121 (1991).Article 

Google Scholar 
Szucs, M., Melbourne, B. A., Tuff, T. & Hufbauer, R. A. The roles of demography and genetics in the early stages of colonization. Proc. R. Soc. B Biol. Sci. 281, 20141073 (2014).Article 

Google Scholar 
Williamson, M. Biological invasions Vol. 15 (Springer, 1996).
Google Scholar 
Dai, Z. C. et al. Synergy among hypotheses in the invasion process of alien plants: A road map within a timeline. Perspect. Plant Ecol. Evol. Syst. 47, 125575 (2020).Article 

Google Scholar 
Briski, E. et al. Beyond propagule pressure: Importance of selection during the transport stage of biological invasions. Front. Ecol. Environ. 16, 345–353 (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Li, Y. & Vitt, D. H. The dynamics of moss establishment: Temporal responses to nutrient gradients. Bryologist 97, 357–364 (1994).Article 

Google Scholar 
Li, Y. & Vitt, D. H. The dynamics of moss establishment: Temporal responses to a moisture gradient. J. Bryol. 18, 677–687 (1995).Article 

Google Scholar 
Wiklund, K. & Rydin, H. Ecophysiological constraints on spore establishment in bryophytes. Funct. Ecol. 18, 907–913 (2004).Article 

Google Scholar 
Zanatta, F. et al. Bryophytes are predicted to lag behind future climate change despite their high dispersal capacities. Nat. Commun. 11, 1–9 (2020).Article 

Google Scholar 
Seaborn, T. J., Goldberg, C. S. & Crespi, E. J. Integration of dispersal data into distribution modeling: What have we done and what have we learned?. Front. Biogeogr. 12, 1–14 (2020).Article 

Google Scholar 
Glime, J. M. Bryophyte Ecology (Vol. 1, Issue Physiological Ecology, Chapter 4–10 Adaptive strategies: vegetative propagules, pp. 1–44). (2021).Guerra, J., Brugués, M., Cano, M. J. & Cros, R. M. Bryum Hedw. in Flora Briofítica Ibérica, Vol. IV, Funariales, Splachnales, Schistostegales, Bryales, Timmiales (eds. Brugués, M. & Cros, R. M.) 105–178 (Universidad de Murcia. Sociedad Española de Briología, 2010).
Google Scholar 
Medina, N. G., Draper, I. & Lara, F. Biogeography of mosses and allies: Does size matter? in Biogeography of microscopic organisms: is everything small everywhere? 209–233 (2011). https://doi.org/10.1017/CBO9780511974878.012Miles, C. J. & Longton, R. E. The role of spores in reproduction in mosses. Bot. J. Linn. Soc. 104, 149–173 (1990).Article 

Google Scholar 
Estébanez, B., Draper, I. & Bujalance, R. M. Bryophytes: An approximation to the simplest land plants. in Biodiversidad. Aproximación a la diversidad botánica y zoológica de España 19 (2011).Frey, W. & Kürschner, H. Asexual reproduction, habitat colonization and habitat maintenance in bryophytes. Flora Morphol. Distrib. Funct. Ecol. Plants 206, 173–184 (2011).Article 

Google Scholar 
Giordano, S. et al. Regeneration from detached leaves of Pleurochaete squarrosa (Brid.) Lindb. in culture and in the wild. J. Bryol. 19, 219–227 (1996).Article 

Google Scholar 
La Farge, C., Williams, K. H. & England, J. H. Regeneration of Little Ice Age bryophytes emerging from a polar glacier with implications of totipotency in extreme environments. Proc. Natl. Acad. Sci. U. S. A. 110, 9839–9844 (2013).Article 
PubMed 
PubMed Central 

Google Scholar 
Robinson, S. C. & Miller, N. G. Bryophyte diversity on Adirondack alpine summits is maintained by dissemination and establishment of vegetative fragments and spores. Bryologist 116, 382–391 (2013).Article 

Google Scholar 
Glime, J. M. Chapter 2–1 Meet the bryophytes. in Bryophyte Ecology 1 (2020).Korpelainen, H., Pohjamo, M. & Laaka-Lindberg, S. How efficiently does bryophyte dispersal lead to gene flow?. J. Hattori Bot. Lab. 205, 195–205 (2005).
Google Scholar 
Schuster, R. M. Phytogeography of the Bryophyta. in New manual of Bryology 1, 463–626 (Hattori Bot. Lab, 1983).Löbel, S., Schröder, B. & Snäll, T. Projected shifts in deadwood bryophyte communities under national climate and forestry scenarios benefit large competitors and impair small species. J. Biogeogr. https://doi.org/10.1111/jbi.14278 (2021).Article 

Google Scholar 
Laaka-Lindberg, S., Korpelainen, H. & Pohjamo, M. Dispersal of asexual propagules in bryophytes. J. Hattori Bot. Lab. 330, 319–330 (2003).
Google Scholar 
Miller, N. G. & Mogensen, G. S. Cyrtomnium hymenophylloides (Bryophyta, Mniaceae) in North America and Greenland: Male plants, sex-differential geographical distribution, and reproductive characteristics. Bryologist 100, 499–506 (1997).Article 

Google Scholar 
Muñoz, J., Felicísimo, Á. M., Cabezas, F., Burgaz, A. R. & Martínez, I. Wind as a long-distance dispersal vehicle in the Southern Hemisphere. Science 304, 1144–1147 (2004).Article 
PubMed 

Google Scholar 
Patiño, J. & Vanderpoorten, A. Bryophyte biogeography. CRC. Crit. Rev. Plant Sci. 37, 175–209 (2018).Article 

Google Scholar 
Pasiche-Lisboa, C. J., Booth, T., Belland, R. J. & Piercey-Normore, M. D. Moss and lichen asexual propagule dispersal may help to maintain the extant community in boreal forests. Ecosphere 10, e02823 (2019).Article 

Google Scholar 
Barbé, M., Fenton, N. J. & Bergeron, Y. So close and yet so far away: Long-distance dispersal events govern bryophyte metacommunity reassembly. J. Ecol. 104, 1707–1719 (2016).Article 

Google Scholar 
Hansson, L., Söderström, L. & Solbreck, C. The ecology of dispersal in relation to conservation. in Ecological principles of nature conservation. Conservation Ecology series: principles, practices and management. (ed. Hansson, L.) (Springer, 1992). https://doi.org/10.1007/978-1-4615-3524-9Chapter 

Google Scholar 
Miller, N. G. & Ambrose, L. J. H. Growth in culture of wind-blown bryophyte gametophyte fragments from Arctic Canada. Bryologist 79, 55 (1976).Article 

Google Scholar 
Barbé, M., Fenton, N. J., Caners, R. & Bergeron, Y. Inter-annual variation in bryophyte dispersal: Linking bryophyte phenophases and weather conditions. Botany 95, 1151–1169 (2017).Article 

Google Scholar 
Chmielewski, M. W. & Eppley, S. M. Forest passerines as a novel dispersal vector of viable bryophyte propagules. Proc. R. Soc. B Biol. Sci. 286, 20182253 (2019).Article 
CAS 

Google Scholar 
Davison, G. W. H. Role of birds in moss dispersal. Br. Birds 69, 65–66 (1976).
Google Scholar 
Heinken, T., Lees, R., Raudnitschka, D. & Runge, S. Epizoochorous dispersal of bryophyte stem fragments by roe deer (Capreolus capreolus) and wild boar (Sus scrofa). J. Bryol. 23, 293–300 (2001).Article 

Google Scholar 
Parsons, J. G. et al. Bryophyte dispersal by flying foxes: A novel discovery. Oecologia 152, 112–114 (2007).Article 
CAS 
PubMed 

Google Scholar 
Glime, J. M. Bryophyte Ecology (Vol. 2, Issue Bryological Interaction) (2021).Ware, C., Bergstrom, D. M., Müller, E. & Alsos, I. G. Humans introduce viable seeds to the Arctic on footwear. Biol. Invasions 14, 567–577 (2012).Article 

Google Scholar 
Shacklette, H. T. Unattached moss polsters on Amchitka Island, Alaska. Bryologist 69, 346–352 (1966).Article 

Google Scholar 
Moles, A. T. & Westoby, M. Seedling survival and seed size: A synthesis of the literature. J. Ecol. 92, 372–383 (2004).Article 

Google Scholar 
Kimmerer, R. W. Patterns of dispersal and establishment of bryophytes colonizing natural and experimental treefall mounds in northern hardwood forests. Bryologist 108, 391–401 (2005).Article 

Google Scholar 
Pérez-Harguindeguy, N. et al. New handbook for standardised measurement of plant functional traits worldwide. Aust. J. Bot. 61, 167–234 (2013).Article 

Google Scholar 
Stieha, C. R., Middleton, A. R., Stieha, J. K., Trott, S. H. & Mcletchie, D. N. The dispersal process of asexual propagules and the contribution to population persistence in Marchantia (Marchantiaceae). Am. J. Bot. 101, 348–356 (2014).Article 
PubMed 

Google Scholar 
Hugonnot, V. Comparative investigations of niche, growth rates and reproduction between the native moss Campylopus pilifer and the invasive C. introflexus. J. Bryol. 39, 79–84 (2017).Article 

Google Scholar 
Benscoter, B. W. Post-fire bryophyte establishment in a continental bog. J. Veg. Sci. 17, 647–652 (2006).Article 

Google Scholar 
Esposito, A., Mazzoleni, S. & Strumia, S. Post-fire bryophyte dynamics in Mediterranean vegetation. J. Veg. Sci. 10, 261–268 (1999).Article 

Google Scholar 
Naeth, M. A. & Wilkinson, S. R. Establishment of restoration trajectories for upland tundra communities on diamond mine wastes in the Canadian arctic. Restor. Ecol. 22, 534–543 (2014).Article 

Google Scholar 
Lamarre, J. J. M. Tundra bryophyte revegetation: novel methods for revegetating northern ecosystems (University of Alberta, 2016).Dierßen, K. Distribution, ecological amplitude and phytosociological characterization of European bryophytes. (Bryophytorum Bibliotheca 56. J. Cramer, Berlin, 289 pp., 2001).Smith, A. J. E. The moss flora of Britain and Ireland (Cambridge University Press, 2004).Book 

Google Scholar 
Casas, C., Brugués, M., Cros, R. M. & Sérgio, C. Handbook of Mosses of the Iberian Peninsula and the Balearic Islands. (Instituts d’Estudis Catalans, 2006).Medina, N., Mazimpaka Nibarere, V., Hortal, J. & Lara García, F. Catálogo de los briófitos epífitos que crecen en bosques de quercíneas del cuadrante noroccidental ibérico. Boletín la Soc. Esp. Briol. 30, 1–30 (2015).
Google Scholar 
Ron Alvarez, M. E. & Vicente, J. Contribución al conocimiento de la flora briológica de Canencia, Sierra de Guadarrama (Madrid). Bot. Complut. https://doi.org/10.5209/BOCM.7415 (1989).Article 

Google Scholar 
Pressel, S., Matcham, H. W. & Duckett, J. G. Studies of protonemal morphogenesis in mosses. XI. Bryum and allied genera: A plethora of propagules. J. Bryol. 29, 241–258 (2007).Article 

Google Scholar 
Söderström, L. & Herben, T. Dynamics of bryophyte metapopulations. in Advances in Briology 6. Population studies (ed. Longton, R. E.) 6, 205–240 (International Association of Briologists. Schweizerbart Science Publishers, 1997).
Google Scholar 
Schneider, C. A., Rasband, W. S. & Eliceiri, K. W. NIH Image to ImageJ: 25 years of image analysis. Nat. Methods 9, 671–675 (2012).Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Cox, E. P. A method of assigning numerical and percentage values to the degree of roundness of sand grains. J. Paleontol. 1, 179–183 (1927).
Google Scholar 
R Core Team. R: A language and environment for Statistical Computing (2021).Kassambara, A. rstatix: Pipe-friendly framework for basic statistical tests (2020).Zeileis, A., Meyer, D. & Hornik, K. Residual-based shadings for visualizing (conditional) independence. J. Comput. Graph. Stat. 16, 507–525 (2007).Article 
MathSciNet 

Google Scholar 
Wickham, H., François, R., Henry, L. & Müller, K. dplyr: A grammar of Data Manipulation (2022).Fox, J. & Weisberg, S. An R Companion to Applied Regression (2019).Maechler, M. et al. robustbase: Basic Robust Statistics (2022).Kassambara, A. ggpubr: ‘ggplot2’ Based Publication Ready Plots (2020).Revelle, W. psych: Procedures for psychological, psychometric, and personality research (2021).Kuhn, M., Jackson, S. & Cimentada, J. corrr: correlations in R. R package version 0.4.3 (2020).Wei, T. & Simko, V. R package ‘corrplot’: visualization of a correlation matrix (Version 0.84) (2017).Wilke, C. O. ggtext: improved text rendering support for ‘ggplot2’ (2020).Auguie, B. gridExtra: miscellaneous functions for ‘Grid’ graphics (2017).Wilke, C. O. cowplot: streamlined plot theme and plot annotations for ‘ggplot2’. R package version 1.1.1 (2020).Stark, L. R., Nichols, L. II., McLetchie, D. N., Smith, S. D. & Zundel, C. Age and sex-specific rates of leaf regeneration in the Mojave Desert moss Syntrichia caninervis. Am. J. Bot. 91, 1–9 (2004).Article 
PubMed 

Google Scholar 
Fernandez-Mendoza, F., Estebanez, B., Gomez-Sanz, D. & Ron, E. Sporophyte-bearing specimens of Pleurochaete squarrosa in Zamora, Spain. Cryptogam. Bryol. 23, 211–215 (2002).
Google Scholar 
Chen, K. H., Liao, H. L., Arnold, A. E., Bonito, G. & Lutzoni, F. RNA-based analyses reveal fungal communities structured by a senescence gradient in the moss Dicranum scoparium and the presence of putative multi-trophic fungi. New Phytol. 218, 1597–1611 (2018).Article 
CAS 
PubMed 

Google Scholar 
Kruijer, H. J. D., Raes, N. & Stech, M. Modelling the distribution of the moss species Hypopterygium tamarisci (Hypopterygiaceae, Bryophyta) in Central and South America. Nov. Hedwigia 91, 399–420 (2010).Article 

Google Scholar 
Van Zanten, B. O. Preliminary report on germination experiments designed to estimate the survival chances of moss spores during aerial trans-oceanic long-range dispersal in the Southern Hemisphere, with particular reference to New Zealand. J. Hattori Bot. Lab. 41, 133–140 (1976).
Google Scholar 
Van Zanten, B. O. Experimental studies on trans-oceanic long-range dispersal of moss spores in the Southern Hemisphere. J. Hattori Bot. Lab. 44, 455–482 (1978).
Google Scholar 
De Meester, L., Gómez, A., Okamura, B. & Schwenk, K. The monopolization hypothesis and the dispersal-gene flow paradox in aquatic organisms. Acta Oecologica 23, 121–135 (2002).Article 

Google Scholar 
Izquieta-Rojano, S. et al. Pleurochaete squarrosa (Brid.) Lindb. as an alternative moss species for biomonitoring surveys of heavy metal, nitrogen deposition and δ15N signatures in a Mediterranean area. Ecol. Indic. 60, 1221–1228 (2016).Article 
CAS 

Google Scholar 
Kimmerer, R. W. & Young, C. C. Effect of gap size and regeneration niche on species coexistence in bryophyte communities. J. Torrey Bot. Soc. 123, 16–24 (1996).Article 

Google Scholar 
Refoyo, P., Peláez, M., García-Rodríguez, M., López-Sánchez, A. & Perea, R. Moss cover and browsing scores as sustainability indicators of mountain ungulate populations in Mediterranean environments. Biodivers. Conserv. https://doi.org/10.1007/s10531-022-02454-1 (2022).Article 

Google Scholar  More

Körner, C. Alpine Plant Life: Functional plant ecology of high mountain ecosystems. (Springer, 2021). https://doi.org/10.1007/978-3-030-59538-8.Pepin, N. et al. Elevation-dependent warming in mountain regions of the world. Nat. Clim. Change 5, 424–430 (2015).Article 

Google Scholar 
Pepin, N. C. et al. Climate changes and their elevational patterns in the mountains of the world. Rev. Geophys. 60, e2020RG000730 (2022).Stewart, I. T. Changes in snowpack and snowmelt runoff for key mountain regions. Hydrol. Process 23, 78–94 (2009).Article 

Google Scholar 
Vorkauf, M., Marty, C., Kahmen, A. & Hiltbrunner, E. Past and future snowmelt trends in the Swiss Alps: the role of temperature and snowpack. Clim. Change 165, 44–62 (2021).Article 

Google Scholar 
Inouye, D. W. Effects of climate change on phenology, frost damage, and floral abundance of montane wildflowers. Ecology 89, 353–362 (2008).Article 
PubMed 

Google Scholar 
Vorkauf, M., Kahmen, A., Körner, C. & Hiltbrunner, E. Flowering phenology in alpine grassland strongly responds to shifts in snowmelt but weakly to summer drought. Alp. Bot. 131, 73–88 (2021).Article 

Google Scholar 
Wipf, S. & Rixen, C. A review of snow manipulation experiments in Arctic and alpine tundra ecosystems. Polar Res. 29, 95–109 (2010).Article 

Google Scholar 
Collins, C. G. et al. Experimental warming differentially affects vegetative and reproductive phenology of tundra plants. Nat. Commun. 12, 3442 (2021).Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Choler, P. Growth response of temperate mountain grasslands to inter-annual variations in snow cover duration. Biogeosciences 12, 3885–3897 (2015).Article 

Google Scholar 
Xie, J. et al. Land surface phenology and greenness in alpine grasslands driven by seasonal snow and meteorological factors. Sci. Total Environ. 725, 138380 (2020).Article 
CAS 
PubMed 

Google Scholar 
Nord, E. A. & Lynch, J. P. Plant phenology: a critical controller of soil resource acquisition. J. Exp. Bot. 60, 1927–1937 (2009).Article 
CAS 
PubMed 

Google Scholar 
Gallinat, A. S., Primack, R. B. & Wagner, D. L. Autumn, the neglected season in climate change research. Trends Ecol. Evol. 30, 169–176 (2015).Article 
PubMed 

Google Scholar 
Rosa, R. K. et al. Plant phenological responses to a long‐term experimental extension of growing season and soil warming in the tussock tundra of Alaska. Glob. Change Biol. 21, 4520–4532 (2015).Article 

Google Scholar 
Livensperger, C. et al. Earlier snowmelt and warming lead to earlier but not necessarily more plant growth. AoB Plants 8, plw021 (2016).Article 
PubMed 
PubMed Central 

Google Scholar 
Körner, C. & Hiltbrunner, E. Why is the alpine flora comparatively robust against climatic warming? Diversity 13, 383–397 (2021).Article 

Google Scholar 
Ma, H. et al. The global distribution and environmental drivers of aboveground versus belowground plant biomass. Nat. Ecol. Evol. 5, 1110–1122 (2021).Article 
PubMed 

Google Scholar 
Iversen, C. M. et al. The unseen iceberg: plant roots in arctic tundra. N. Phytol. 205, 34–58 (2015).Article 

Google Scholar 
Abramoff, R. Z. & Finzi, A. C. Are above‐ and below‐ground phenology in sync? N. Phytol. 205, 1054–1061 (2015).Article 

Google Scholar 
Liu, H. et al. Phenological mismatches between above- and belowground plant responses to climate warming. Nat. Clim. Change 12, 97–102 (2022).Article 
CAS 

Google Scholar 
Rixen, C. et al. Winters are changing: snow effects on Arctic and alpine tundra ecosystems. Arct. Sci. 1–37 (2022) https://doi.org/10.1139/as-2020-0058.Johnson, M. G., Tingey, D. T., Phillips, D. L. & Storm, M. J. Advancing fine root research with minirhizotrons. Environ. Exp. Bot. 45, 263–289 (2001).Article 
PubMed 

Google Scholar 
Atkinson, J. A., Pound, M. P., Bennett, M. J. & Wells, D. M. Uncovering the hidden half of plants using new advances in root phenotyping. Curr. Opin. Biotech. 55, 1–8 (2019).Article 
CAS 
PubMed 

Google Scholar 
Radville, L., McCormack, M. L., Post, E. & Eissenstat, D. M. Root phenology in a changing climate. J. Exp. Bot. 67, 3617–3628 (2016).Article 
CAS 
PubMed 

Google Scholar 
Blume-Werry, G. The belowground growing season. Nat. Clim. Change 12, 11–12 (2022).Article 

Google Scholar 
Wipf, S., Stoeckli, V. & Bebi, P. Winter climate change in alpine tundra: plant responses to changes in snow depth and snowmelt timing. Clim. Change 94, 105–121 (2009).Article 

Google Scholar 
Baptist, F., Flahaut, C., Streb, P. & Choler, P. No increase in alpine snowbed productivity in response to experimental lengthening of the growing season. Plant Biol. 12, 755–764 (2010).Article 
CAS 
PubMed 

Google Scholar 
Vitasse, Y. et al. ‘Hearing’ alpine plants growing after snowmelt: ultrasonic snow sensors provide long-term series of alpine plant phenology. Int J. Biometeorol. 61, 349–361 (2017).Article 
PubMed 

Google Scholar 
Blume‐Werry, G., Jansson, R. & Milbau, A. Root phenology unresponsive to earlier snowmelt despite advanced above‐ground phenology in two subarctic plant communities. Funct. Ecol. 31, 1493–1502 (2017).Article 

Google Scholar 
Darrouzet‐Nardi, A. et al. Limited effects of early snowmelt on plants, decomposers, and soil nutrients in Arctic tundra soils. Ecol. Evol. 9, 1820–1844 (2019).Article 
PubMed 
PubMed Central 

Google Scholar 
Ernakovich, J. G. et al. Predicted responses of arctic and alpine ecosystems to altered seasonality under climate change. Glob. Change Biol. 20, 3256–3269 (2014).Article 

Google Scholar 
Keller, F. & Körner, C. The role of photoperiodism in alpine plant development. Arct. Antarct. Alp. Res 35, 361–368 (2003).Article 

Google Scholar 
Hiltbrunner, E., Arnaiz, J. & Körner, C. Biomass allocation and seasonal non-structural carbohydrate dynamics do not explain the success of tall forbs in short alpine grassland. Oecologia 1–15 (2021) https://doi.org/10.1007/s00442-021-04950-7.Inauen, N., Körner, C. & Hiltbrunner, E. No growth stimulation by CO2 enrichment in alpine glacier forefield plants. Glob. Change Biol. 18, 985–999 (2012).Article 

Google Scholar 
Möhl, P., Hiltbrunner, E. & Körner, C. Halving sunlight reveals no carbon limitation of aboveground biomass production in alpine grassland. Glob. Change Biol. 26, 1857–1872 (2020).Article 

Google Scholar 
Porter, J. R. & Gawith, M. Temperatures and the growth and development of wheat: a review. Eur. J. Agron. 10, 23–36 (1999).Article 

Google Scholar 
Parent, B., Turc, O., Gibon, Y., Stitt, M. & Tardieu, F. Modelling temperature-compensated physiological rates, based on the co-ordination of responses to temperature of developmental processes. J. Exp. Bot. 61, 2057–2069 (2010).Article 
CAS 
PubMed 

Google Scholar 
Körner, C. H. & Woodward, F. I. The dynamics of leaf extension in plants with diverse altitudinal ranges. Oecologia 72, 279–283 (1987).Article 
PubMed 

Google Scholar 
Nagelmüller, S., Hiltbrunner, E. & Körner, C. Low temperature limits for root growth in alpine species are set by cell differentiation. AoB Plants 9, plx054 (2017).Article 
PubMed 
PubMed Central 

Google Scholar 
Starr, G., Oberbauer, S. F. & Pop, E. W. Effects of lengthened growing season and soil warming on the phenology and physiology of Polygonum bistorta. Glob. Change Biol. 6, 357–369 (2000).Article 

Google Scholar 
Yoshie, F. Vegetative phenology of alpine plants at Tateyama Murodo-daira in central Japan. J. Plant Res. 123, 675–688 (2010).Article 
PubMed 

Google Scholar 
Jonas, T., Rixen, C., Sturm, M. & Stoeckli, V. How alpine plant growth is linked to snow cover and climate variability. J. Geophys. Res. 113, G03013 (2008).
Google Scholar 
Wang, H. et al. Alpine grassland plants grow earlier and faster but biomass remains unchanged over 35 years of climate change. Ecol. Lett. 23, 701–710 (2020).Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Frei, E. R. & Henry, G. H. R. Long-term effects of snowmelt timing and climate warming on phenology, growth, and reproductive effort of Arctic tundra plant species. Arct. Sci. 1–22 (2021) https://doi.org/10.1139/as-2021-0028.Schäppi, B. & Körner, C. Growth responses of an alpine grassland to elevated CO2. Oecologia 105, 43–52 (1996).Article 
PubMed 

Google Scholar 
Aloni, R. Role of hormones in controlling vascular differentiation and the mechanism of lateral root initiation. Planta 238, 819–830 (2013).Article 
CAS 
PubMed 

Google Scholar 
Sloan, V. L., Fletcher, B. J. & Phoenix, G. K. Contrasting synchrony in root and leaf phenology across multiple sub‐Arctic plant communities. J. Ecol. 104, 239–248 (2016).Article 
CAS 

Google Scholar 
Nagelmüller, S., Hiltbrunner, E. & Körner, C. Critically low soil temperatures for root growth and root morphology in three alpine plant species. Alp. Bot. 126, 11–21 (2016).Article 

Google Scholar 
Woo, H. R., Kim, H. J., Lim, P. O. & Nam, H. G. Leaf senescence: systems and dynamics aspects. Annu. Rev. Plant Biol. 70, 1–30 (2019).Article 

Google Scholar 
Liu, Z., Marella, C. B. N., Hartmann, A., Hajirezaei, M. R. & Wirén, Nvon An age-dependent sequence of physiological processes defines developmental root senescence. Plant Physiol. 181, 993–1007 (2019).Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Ryser, P., Puig, S., Müller, M. & Munné-Bosch, S. Abscisic acid responses match the different patterns of autumn senescence in roots and leaves of Iris versicolor and Sparganium emersum. Environ. Exp. Bot. 176, 104097 (2020).Article 
CAS 

Google Scholar 
Budge, K., Leifeld, J., Hiltbrunner, E. & Fuhrer, J. Alpine grassland soils contain large proportion of labile carbon but indicate long turnover times. Biogeosciences 8, 1911–1923 (2011).Article 
CAS 

Google Scholar 
Solly, E. F. et al. Unravelling the age of fine roots of temperate and boreal forests. Nat. Commun. 9, 3006 (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Trumbore, S. E., Sierra, C. A. & Pries, C. E. H. Radiocarbon and climate change, mechanisms, applications and laboratory techniques. 45–82 (2016) https://doi.org/10.1007/978-3-319-25643-6_3.Windmaißer, T. & Reisch, C. Long-term study of an alpine grassland: local constancy in times of global change. Alp. Bot. 123, 1–6 (2013).Article 

Google Scholar 
De Witte, L. C. D., Armbruster, G. F. J., Gielly, L., Taberlet, P. & Stöcklin, J. AFLP markers reveal high clonal diversity and extreme longevity in four key arctic‐alpine species. Mol. Ecol. 21, 1081–1097 (2012).Article 
PubMed 

Google Scholar 
Landolt, E. Unsere Alpenflora. (SAC-Verlag, 2012).Puşcaş, M. & Choler, P. A biogeographic delineation of the European Alpine System based on a cluster analysis of Carex curvula-dominated grasslands. Flora – Morphol. Distrib. Funct. Ecol. Plants 207, 168–178 (2012).Article 

Google Scholar 
Grabherr, G., Mahr, E. & Reisigl, H. Nettoprimärproduktion und Reproduktion in einem Krummseggenrasen (Caricetum curvulae) der Otztaler Alpen, Tirol. Oecologia Plant. 13, 227–251 (1978).
Google Scholar 
Chiang, C., Bånkestad, D. & Hoch, G. Reaching natural growth: light quality effects on plant performance in indoor growth facilities. Plants 9, 1273 (2020).Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Chiang, C., Olsen, J. E., Basler, D., Bånkestad, D. & Hoch, G. Latitude and weather influences on sun light quality and the relationship to tree growth. Forests 10, 610–621 (2019).Article 

Google Scholar 
Richardson, A. D. et al. Use of digital webcam images to track spring green-up in a deciduous broadleaf forest. Oecologia 152, 323–334 (2007).Article 
PubMed 

Google Scholar 
Jiang, Y. & Li, C. Convolutional neural networks for image-based high-throughput plant phenotyping: a review. Plant Phenomics 2020, 4152816 (2020).Article 
PubMed 
PubMed Central 

Google Scholar 
Möller, B. et al. rhizoTrak: a flexible open source Fiji plugin for user-friendly manual annotation of time-series images from minirhizotrons. Plant Soil 444, 519–534 (2019).Article 

Google Scholar 
Schindelin, J. et al. Fiji: an open-source platform for biological-image analysis. Nat. Methods 9, 676–682 (2012).Article 
CAS 
PubMed 

Google Scholar 
Smith, A. G., Petersen, J., Selvan, R. & Rasmussen, C. R. Segmentation of roots in soil with U-Net. Plant Methods 16, 13–27 (2020).Article 
PubMed 
PubMed Central 

Google Scholar 
Seethepalli, A. et al. Rhizovision crown: an integrated hardware and software platform for root crown phenotyping. Plant Phenomics 2020, 3074916 (2020).Article 
PubMed 
PubMed Central 

Google Scholar 
R Core Team. R: A language and environment for statistical computing. (R Foundation for Statistical Computing, 2021).Wood, S. N. Fast stable restricted maximum likelihood and marginal likelihood estimation of semiparametric generalized linear models. J. R. Stat. Soc. Ser. B Stat. Methodol. 73, 3–36 (2011).Article 
MathSciNet 
MATH 

Google Scholar 
Lenth, R. V. emmeans: Estimated Marginal Means, aka Least-Squares Means. R package version 1.6.2-1. (2021).Möhl P., von Büren R. S. & Hiltbrunner E. Data from: Growth of alpine grassland will start and stop earlier under climate warming figshare. https://doi.org/10.6084/m9.figshare.20440497 (2022). More