Benthic communities at two remote Pacific coral reefs: effects of reef habitat, depth, and wave energy gradients on spatial patterns.

Kingman Reef and Palmyra Atoll in the central Pacific are among the most remote coral reefs on the planet. Here we describe spatial patterns in their benthic communities across reef habitats and depths, and consider these in the context of oceanographic gradients. Benthic communities at both locations were dominated by calcifying organisms (54-86% cover), namely hard corals (20-74%) and crustose coralline algae (CCA) (10-36%). While turf algae were relatively common at both locations (8-22%), larger fleshy macroalgae were virtually absent at Kingman (<1%) and rare at Palmyra (0.7-9.3%). Hard coral cover was higher, but with low diversity, in more sheltered habitats such as Palmyra's backreef and Kingman's patch reefs. Almost exclusive dominance by slow-growing Porites on Kingman's patch reefs provides indirect evidence of competitive exclusion, probably late in a successional sequence. In contrast, the more exposed forereef habitats at both Kingman and Palmyra had higher coral diversity and were characterized by fast-growing corals (e.g., Acropora and Pocillopora), indicative of more dynamic environments. In general at both locations, soft coral cover increased with depth, likely reflecting increasingly efficient heterotrophic abilities. CCA and fleshy macroalgae cover decreased with depth, likely due to reduced light. Cover of other calcified macroalgae, predominantly Halimeda, increased with depth. This likely reflects the ability of many calcifying macroalgae to efficiently harvest light at deeper depths, in combination with an increased nutrient supply from upwelling promoting growth. At Palmyra, patterns of hard coral cover with depth were inconsistent, but cover peaked at mid-depths at Kingman. On Kingman's forereef, benthic community composition was strongly related to wave energy, with hard coral cover decreasing and becoming more spatially clustered with increased wave energy, likely as a result of physical damage leading to patches of coral in localized shelter. In contrast, the cover of turf algae at Kingman was positively related to wave energy, reflecting their ability to rapidly colonize newly available space. No significant patterns with wave energy were observed on Palmyra's forereef, suggesting that a more detailed model is required to study biophysical coupling there. Kingman, Palmyra, and other remote oceanic reefs provide interesting case studies to explore biophysical influences on benthic ecology and dynamics.