Invertebrate trophic structure on marine ferromanganese and phosphorite hardgrounds

The Southern California Borderland hosts a variety of geologic and oceanographic features that allow for diverse habitats to occur in a restricted region with a strong oxygen minimum zone (OMZ) and hard substrates. These include ferromanganese (FeMn) crusts and phosphorites targeted for deep-seabed mining in other regions. Baseline studies regarding hardground macro- (> 0.3 mm) and megafaunal (> 2 cm) invertebrates are lacking, although they contribute to understanding nutrient cycling and resilience of deep-sea communities to ocean deoxygenation, fishing, or mineral extraction. With the goal of understanding how substrate type, depth, and dissolved oxygen concentration influence invertebrate trophic structure, we surveyed delta C-13 and delta N-15 values of invertebrates on hard substrates on the Southern California Borderland margin along a depth gradient (120-2400 m) through the OMZ at inshore (< 100 km from shore) and offshore (100-250 km from shore) sites, using generalized additive models and community-level metrics. Macrofaunal isotopic values correlate with substrate type, exhibiting higher trophic diversity on FeMn crusts and specialized communities on phosphorites. Megafaunal isotopic values correlate with proximity to shore; animals offshore seem to depend more on phytoplanktonic production than animals inshore. In general, delta N-15 increased with decreasing dissolved oxygen and increasing depth, possibly due to remineralization processes within the OMZ and with depth. We discuss how feeding modes and community composition might influence the observed patterns. This study elucidates the importance of the environmental context in shaping invertebrate trophic structure on continental margins and provides baseline knowledge that may be useful in regions where these minerals are targeted for extraction.