Coral reefs have changed radically in the last few decades with reefs in the Caribbean now averaging 13% coral cover and 40% macroalgal cover (mostly Dictyota and Lobophora). So, it is time we re-evaluate which species are key to the process of herbivory in these new conditions. The role herbivorous fishes play in controlling macroalgae is often considered by managers and researchers at a guild or family level, but greater resolution is needed to understand the impact of herbivores more fully. We performed feeding assays and behavioral observations of fish feeding to quantify the removal of the most common macroalgae by different herbivorous fish species. In total, we ran 34 h-long trials using Dictyota and Lobophora across two sites and conducted over 34 h of observation of 105 fish from eight species in the Cayman Islands, Caribbean. We show that many nominal herbivores did not consume macroalgae but instead targeted the epibionts on macroalgae and other substrates. In fact, only three fish taxa consumed macroalgae as a significant proportion of their feeding: one species of surgeonfish (Acanthurus coeruleus), one species of parrotfish (Sparisoma aurofrenatum), and the third, the chubs (Kyphosus spp.), is a group of species which is not consistently considered as part of the herbivore community in the Caribbean. From our observations, an individual A. coeruleus can consume ∼44 g of Dictyota per day, while S. aurofrenatum can consume ∼50 g and Kyphosus spp. can consume ∼100 g. These values are significantly more than all other herbivorous fish species and suggest these three taxa are key macroalgal consumers in the Caribbean. These results highlight that disentangling the role of individual herbivore species is necessary for critical species to be identified and protected. Furthermore, as reef conditions change, we need to reevaluate the key functions and species to be more effective at protecting and managing these important ecosystems. With far higher macroalgal coverage than in the past, the few browsing species that remove macroalgae may be increasingly important in promoting reef health.

Many reefs have shifted from coral and fish dominated habitats to less productive macroalgal dominated habitats, and current research is investigating means of reversing this phase shift. In the tropical Pacific, overfished reefs with inadequate herbivory can become dominated by the brown alga Sargassum polycystum. This alga suppresses recruitment and survival of corals and fishes, thus limiting the potential for reef recovery. Here we investigate the mechanisms that reinforce S. polycystum dominance and show that in addition to negatively affecting other species, this species acts in a self-reinforcing manner, positively promoting survival and growth of conspecifics. We found that survival and growth of both recruit-sized and mature S. polycystum fronds were higher within Sargassum beds than outside the beds and these results were found in both protected and fished reefs. Much of this benefit resulted from reduced herbivory within the Sargassum beds, but adult fronds also grew ~50% more within the beds even when herbivory did not appear to be occurring, suggesting some physiological advantage despite the intraspecific crowding. Thus via positive feedbacks, S. polycystum enhances its own growth and resistance to herbivores, facilitating its dominance (perhaps also expansion) and thus its resilience on degraded reefs. This may be a key feedback mechanism suppressing the recovery of coral communities in reefs dominated by macroalgal beds.

Marine Protected Areas (MPAs) are often established to mitigate the effects of overfishing and other human disturbances. In Fiji these are locally managed and, where enforced, have significantly higher coral cover, higher fish biomass, and lower seaweed cover than in the adjacent, unprotected reefs (non-MPAs). We investigated how the isotopic signatures of a common, mid-level consumer, Epinephelus merra, differed among three small (0.5- 0.8km2) MPAs versus adjacent, unprotected reefs. Isotopic ratios suggested that the fish in the MPAs fed higher in the food chain than those in the adjacent non-MPAs, despite being slightly smaller in size. Calculations using a brown alga as representative of the basal level of the food chain estimate this difference to be about half a trophic level. Thus, the isotopic ratio of a mid-level consumer can be noticeably altered over scales of only a few hundred meters. This may result from more complete food webs and hence greater prey choice and availability in the MPAs and implies that MPAs affect not only species’ abundance and diversity, but also diet composition and trophic biology of member individuals. Our findings suggest E. merra exhibits considerable site fidelity in its feeding biology and thus provides a localized isotopic signal of its reef of residence. If the isotopic signal of this mid-level carnivore is reflective of the composition of the food web beneath it, the signal might provide an easily obtained indication of reef conditions in that area.

There are many examples of macroalgae inducing defence in response to small invertebrate herbivores like amphipods, isopods, and gastropods but few cases of induction in response to vertebrate macrograzers like herbivorous fishes. This may be because larger grazers rapidly consume large quantities of seaweed before induction can occur, thus selecting for constitutive rather than induced defences. Alternatively, the pattern could occur because induction due to feeding by macrograzers is less commonly investigated. In Fiji, field assays with the brown macroalga Sargassum polycystum demonstrated that thalli growing in marine protected areas (MPAs) with abundant herbivorous fishes were significantly less palatable than those growing in adjacent fished areas (non-MPAs) with few herbivorous fishes. This significant preference occurred in 11 of 13 trials over 5 time periods and across 3 pairs of MPAs and spatially associated non-MPAs. This preference was not positively associated with algal nitrogen content or with the toughness of algal fronds. When S. polycystum ramets were taken from the non-MPA and half were partially grazed by fishes while the other half were protected from grazing, new growth from the controls was strongly preferred to new growth from the previously grazed ramets although these fronds originated from the same holdfast. This suggests that S. polycystum upregulates defences (probably chemical) in response to grazing by herbivorous fishes. This is one of the few published examples of induction of macroalgal defence in response to feeding by large, mobile grazers. It is unclear whether induced defences against fishes are rare or just under-investigated.

Many Caribbean reefs have shifted from coral dominance to macroalgal dominance, often by brown algae such as Dictyota and Lobophora. However, the north side of Cayman Brac in the Cayman Islands is dominated seasonally by the green macroalga Microdictyon (percent cover of Microdictyon is 4% in the winter and ~40% of the reef in the summer), although it is absent from the south side of the island and the remainder of the country. Indeed, Microdictyon is rare in much of the Caribbean, so this situation on Cayman Brac provides an opportunity to investigate the conditions that facilitate its distribution and dominance. The impact of herbivory, competition, nutrient input and other abiotic conditions were examined as factors that could influence the distribution and dominance of Microdictyon. While herbivory or nutrient input are frequently found to be key drivers of benthic community composition on coral reefs, here consumption of Microdictyon by herbivores was low, and thus, the alga was not subject to strong top-down control by herbivory. So, in this case, neither herbivore abundance nor feeding preference appeared to influence the distribution of Microdictyon. Nutrient input was also similar to both sides of the island suggesting nutrients played little role in differential distribution. But, in a controlled transplant experiment where Microdictyon was protected from herbivory and competition, it experienced almost complete mortality (93.3%) when transplanted to the south side, compared to only 11.8% mortality on the north. The south side was exposed to the strongest wave action 92% of the days in our study and was on average a slight, but significant 0.2°C warmer. Thus, these data suggest physical forcing (i.e. wave exposure) is the most likely factor dictating Microdictyon distribution. Conversely, a combination of reduced herbivory and increased competitive strength may explain the seasonal dominance of Microdictyon on the north side of Cayman Brac. Microdictyon was a competitive inferior to the other common algae in winter but increased in competitive strength to equal other species in summer. These results add to the literature on Microdictyon and on the forces impacting benthic community structure of coral reefs.