Benthic modification and biotic associations at natural and artificial habitats excavated by Epinephelus morio and Lutjanus campechanus

Pockmarks are abundant seafloor features worldwide and, in West Florida Shelf waters <110 m deep, are thought to be sites of sediment excavation primarily by red grouper Epinephelus morio , although red snapper Lutjanus campechanus also excavate sediment. During 2014-2017, side-scan sonar (445 kHz) was used to locate and deploy stereo-baited remote underwater video arrays within view of 202 such excavations in waters 17-110 m deep on the West Florida Shelf off the Florida Panhandle and Peninsula. Three excavation habitat classes included 73 isolated excavations on open sand, 74 associated with low-relief hard bottom, and 55 associated with artificial reefs. Physical characteristics of excavations varied between regions, among habitats, and with depth; mean diameter (±1 SE) was 9.9 ± 0.3 m (range: 3-24.6 m). Excavations not around artificial reefs contained 6.9 ± 0.5 m 2 (0-27.7 m 2 ) of exposed rock, and epibenthic growth covered 33 ± 2% of the interiors. Members of 99 fish genera were identified. Fish abundance was greatest at isolated excavations which showed similar evenness to excavated artificial reefs; diversity was higher at excavated low-relief hard bottom. L. campechanus was much more common in Panhandle waters, especially at excavated artificial reefs which had subsided 0.8 ± 0.1 vertical meters below the seafloor (i.e. 48 ± 4% of the structure). These biotic and abiotic characteristics of excavations highlight the importance of E. morio ’s ecosystem-engineering services and provide new insight into the contributions of L. campechanus in creating or maintaining excavations at natural and anthropogenic habitats.