An Integrative Ecological Drought Framework to Span Plant Stress to Ecosystem Transformation

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An Integrative Ecological Drought Framework to Span Plant Stress to Ecosystem Transformation Seth M. Munson,1* John B. Bradford,1 and Kevin R. Hultine2 1

U.S. Geological Survey, Southwest Biological Science Center, Flagstaff, Arizona 86001, USA; 2Department of Research, Conservation, and Collections, Desert Botanical Garden, Phoenix, Arizona 85008, USA

ABSTRACT Droughts have increased globally in the twentyfirst century and are expected to become more extreme and widespread in the future. Assessments of how drought affects plants and ecosystems lack consistency in scope and methodology, confounding efforts to mechanistically interpret structural and functional impacts and predict future transformations under climate change. To promote integration among studies, we identify water deficit conditions that are ecologically meaningful, clarify the stages in which ecological drought progresses, and consider approaches to synthesize drought effects across multiple species and ecosystems. This improved ecological drought framework reveals advantages of using different ecological drought metrics and strengthens approaches to distinguish ecosystem stress from crossing an irreversible

threshold. We employ several well-studied examples from water-limited ecosystems, which contain plants that are often at their physiological limits and highly responsive to climate variability. We suggest that emerging research on early warning signs, drought recovery, and the effects of land management interventions be incorporated into the ecological drought framework. An integrative approach to understand ecological drought can accelerate scientific advancement and create opportunity to adapt and prepare for crossing irreversible ecosystem thresholds. Key words: Climate change and extremes; Critical threshold, Plant and ecosystem sensitivity; Land management; Resistance and resilience; Tipping point; Water availability.

HIGHLIGHTS  An integrative ecological drought framework advances prediction of ecosystem response  Differentiation among ecosystem resistance, stress, and threshold responses is essential  Drought recovery and adaptation strategies are ripe for new research Received 7 February 2020; accepted 28 August 2020 Author Contributions SMM and JBB conceived the review. SMM, JBB, and KRH wrote the paper. *Corresponding author; e-mail: [email protected]

S. M. Munson and others

INTRODUCTION An increase in the frequency and severity of drought has emerged as a trend in the early twenty-first century and is projected to intensify with warming temperatures (IPCC 2014). The consequences of water deficit on the performance of individual organisms to ecosystems have become widely recognized (Allen and others 2010; Zhao and Running 2010; Munson and others 2013; Anderegg and others 2016a). Although the effects of meteorological or climatological drought—a period of abnormal precipitation deficit that causes a hydrologic imbalance (AMS 2013; IPCC 2014)—are relatively well defined for agriculture, surface and subsurface water supply, and h