Jennifer L. Ruesink ^1* Katie Houle ² Kristy Kroeker ³ Brett R. Dumbauld ⁴ Fiona C. Boardman ¹ Nathaniel S. Lewis ⁵ Brooke A. McIntyre ⁶ Andrew D. Suhrbier ² Bobbi Hudson ²

• ¹ University of Washington, Seattle, United States
• ² Pacific Shellfish Institute, Olympia, Washington, United States
• ³ University of California, Santa Cruz, Santa Cruz, California, United States
• ⁴ United States Department of Agriculture (USDA), Washington, District of Columbia, United States
• ⁵ Oregon State University, Corvallis, Oregon, United States
• ⁶ Agricultural Research Service, United States Department of Agriculture, Washington D.C., District of Columbia, United States

Species persist through resistance and recovery traits induced by natural stressors. Whether human activities exacerbate or moderate the effects of natural stressors is an open question. Because eelgrass (Zostera marina) creates coastal habitat, its response to natural stressors and human activities is of particular management importance. In this study, traits of intertidal eelgrass were examined across cumulative stressors of emersion and oyster culture, including two culture types: oysters grown directly on sediment (ground culture) or supported by gear above the sediment (off-bottom culture). Summer eelgrass was larger above-and below-ground and branched less than in spring, while density and cover were similar seasonally but declined at higher elevation and in ground culture. Eelgrass traits were divided into those related to resistance (larger above-ground size and below-ground storage) and recovery (increased flowering, branching, and rhizome extension). Resistance traits responded additively to intertidal elevation and oyster culture, with above-and below-ground size reduced in all conditions except that rhizome mass was maintained in ground culture. Smaller above-ground size may confer resistance to emersion stress, as a departure from expectations for other stressors. For resilience traits, flowering increased and internode length declined at higher elevation, whereas these traits did not change in ground culture, and off-bottom culture was associated with shorter internodes (additive cumulative stressor) and tended to moderate the enhanced flowering at higher elevations (non-additive, P=0.058). Transitory disturbance in ground culture may reduce eelgrass density with few effects on resilience, whereas offbottom culture involves longer-term gear placement and trait responses by eelgrass.