Governments, businesses, and lenders worldwide are adopting an objective of no net loss (NNL) of biodiversity that is often partly achieved through biodiversity offsetting within a hierarchy of mitigation actions. Offsets aim to balance residual losses of biodiversity caused by development in one location with commensurate gains at another. Although ecological challenges to achieve NNL are debated, the associated gains and losses for local stakeholders have received less attention. International best practice calls for offsets to make people no worse off than before implementation of the project, but there is a lack of clarity concerning how to achieve this with regard to people's use and nonuse values for biodiversity, especially given the inevitable trade-offs when compensating biodiversity losses with gains elsewhere. This is particularly challenging for countries where poor people depend on natural resources. Badly planned offsets can exacerbate poverty, and development and offset impacts can vary across spatial-temporal scales and by location, gender, and livelihood. We conceptualize the no-worse-off principle in the context of NNL of biodiversity, by exploring for whom and how the principle can be achieved. Changes in the spatial and temporal distribution of biodiversity-related social impacts of a development and its associated offset can lead to social inequity and negatively impact people's well-being. The level of aggregation (regional, village, interest group, household, and individual) at which these social impacts are measured and balanced can again exacerbate inequity in a system. We propose that a determination that people are no worse off, and preferably better off, after a development and biodiversity offset project than they were before the project should be based on the perceptions of project-affected people (assessed at an appropriate level of aggregation); that their well-being associated with biodiversity losses and gains should be at least as good as it was before the project; and that this level of well-being should be maintained throughout the project life cycle. Employing this principle could help ensure people are no worse off as a result of interventions to achieve biodiversity NNL.

Understanding adaptation by natural selection requires understanding the genetic factors that determine which beneficial mutations are available for selection. Here, using experimental evolution of rifampicinresistant Pseudomonas aeruginosa, we show that different genotypes vary in their capacity for adaptation to the cost of antibiotic resistance. We then use sequence data to show that the beneficial mutations associated with fitness recovery were specific to particular genetic backgrounds, suggesting that genotypes had access to different sets of beneficial mutations. When we manipulated the supply rate of beneficial mutations, by altering effective population size during evolution, we found that it constrained adaptation in some selection lines by restricting access to rare beneficial mutations, but that the effect varied among the genotypes in our experiment. These results suggest that mutational neighbourhood varies even among genotypes that differ by a single amino acid change, and this determines their capacity for adaptation as well as the influence of population biology processes that alter mutation supply rate.

Loss of habitats or ecosystems arising from development projects (e.g., infrastructure, resource extraction, urban expansion) are frequently addressed through biodiversity offsetting. As currently implemented, offsetting typically requires an outcome of "no net loss" of biodiversity, but only relative to a baseline trajectory of biodiversity decline. This type of "relative" no net loss entrenches ongoing biodiversity loss, and is misaligned with biodiversity targets that require "absolute" no net loss or "net gain." Here, we review the limitations of biodiversity offsetting, and in response, This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

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Many nations use ecological compensation policies to address negative impacts of development projects and achieve No Net Loss (NNL) of biodiversity and ecosystem services. Yet, failures are widely reported. We use spatial simulation models to quantify potential net impacts of alternative compensation policies on biodiversity (indicated by native vegetation) and two ecosystem services (carbon storage, sediment retention) across four case studies (in Australia, Brazil, Indonesia, Mozambique). No policy achieves NNL of biodiversity in any case study. Two factors limit their potential success: the land available for compensation (existing vegetation to protect or cleared land to restore), and expected counterfactual biodiversity losses (unregulated vegetation clearing). Compensation also fails to slow regional biodiversity declines because policies regulate only a subset of sectors, and expanding policy scope requires more land than is available for compensation activities. Avoidance of impacts remains essential in achieving NNL goals, particularly once opportunities for compensation are exhausted.