This is an author produced version of a paper published in Nature. This paper has been peer-reviewed and is proof-corrected, but does not include the journal pagination. The most unique feature of Earth is the existence of life, and the most extraordinary feature of life is its diversity. Approximately 9 million types of plants, animals, protists and fungi inhabit the earth. So, too, do 7 billion people. Two decades ago, at the first Earth Summit, the vast majority of the world's nations declared that human actions were dismantling Earth's ecosystems, eliminating genes, species, and biological traits at an alarming rate. This observation led to a daunting question: How 30 will loss of biological diversity alter the functioning of ecosystems and their ability to provide society with the goods and services needed to prosper?

The Millennium Ecosystem Assessment (MA) introduced a new framework for analyzing social-ecological systems that has had wide influence in the policy and scientific communities. Studies after the MA are taking up new challenges in the basic science needed to assess, project, and manage flows of ecosystem services and effects on human well-being. Yet, our ability to draw general conclusions remains limited by focus on discipline-bound sectors of the full social-ecological system. At the same time, some polices and practices intended to improve ecosystem services and human well-being are based on untested assumptions and sparse information. The people who are affected and those who provide resources are increasingly asking for evidence that interventions improve ecosystem services and human well-being. New research is needed that considers the full ensemble of processes and feedbacks, for a range of biophysical and social systems, to better understand and manage the dynamics of the relationship between humans and the ecosystems on which they rely. Such research will expand the capacity to address fundamental questions about complex socialecological systems while evaluating assumptions of policies and practices intended to advance human well-being through improved ecosystem services.

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The science and management of infectious disease are entering a new stage. Increasingly public policy to manage epidemics focuses on motivating people, through social distancing policies, to alter their behavior to reduce contacts and reduce public disease risk. Person-to-person contacts drive human disease dynamics. People value such contacts and are willing to accept some disease risk to gain contact-related benefits. The cost-benefit trade-offs that shape contact behavior, and hence the course of epidemics, are often only implicitly incorporated in epidemiological models. This approach creates difficulty in parsing out the effects of adaptive behavior. We use an epidemiological-economic model of disease dynamics to explicitly model the trade-offs that drive person-toperson contact decisions. Results indicate that including adaptive human behavior significantly changes the predicted course of epidemics and that this inclusion has implications for parameter estimation and interpretation and for the development of social distancing policies. Acknowledging adaptive behavior requires a shift in thinking about epidemiological processes and parameters.susceptible-infected-recovered model | R 0 | reproductive number | bioeconomics T he science and management of infectious disease is entering a new stage. The increasing focus on incentive structures to motivate people to engage in social distancing-reducing interpersonal contacts and hence public disease risk (1)-changes what health authorities need from epidemiological models. Social distancing is not new-for centuries humans quarantined infected individuals and shunned the obviously ill, but new approaches are being used to deal with modern social interactions. Scientific development of social distancing public policies requires that epidemiological models explicitly address behavioral responses to disease risk and other incentives affecting contact behavior. This paper models the role of adaptive behavior in an epidemiological system. Recognizing adaptive behavior means explicitly incorporating behavioral responses to disease risk and other incentives into epidemiological models (2, 3). The workhorse of modern epidemiology, the compartmental epidemiological model (4, 5), does not explicitly include behavioral responses to disease risk. The transmission factors in these models combine and confound human behavior and biological processes. We develop a simple compartmental model that explicitly incorporates adaptive behavior and show that this modification alters understanding of standard epidemiological metrics. For example, the basic reproductive number, R 0 , is a function of biological processes and human behavior, but R 0 lacks a behavioral interpretation in the existing literature. Biological and behavioral feedbacks muddle R 0 's biological interpretation and confound its estimation.Prior approaches that incorporate behavior into epidemiological models generally fall into three categories: specification of nonlinear contact rate functions, expanded epidemiologi...