Evidence for managing herbivores for reef resilience.

Herbivore management is an important tool for resilience-based approaches to coral reef conservation, and evidence-based science is needed to enact successful management. We synthesized data from multiple monitoring programs in Hawai'i to measure herbivore biomass and benthic condition over a 10-year period preceding any major coral bleaching. We analysed data from 20 242 transects alongside data on 27 biophysical and human drivers and found herbivore biomass was highly variable throughout Hawai'i, with high values in remote locations and the lowest values near population centres. Both human and biophysical drivers explained variation in herbivore biomass, and among the human drivers both fishing and land-based pollution had negative effects on biomass. We also found evidence that herbivore functional group biomass is strongly linked to benthic condition, and that benthic condition is sensitive to changes in herbivore biomass associated with fishing. We show that when herbivore biomass is below 80% of potential biomass, benthic condition is predicted to decline. We also show that a range of management actions, including area-specific fisheries regulations and gear restrictions, can increase parrotfish biomass. Together, these results provide lines of evidence to support managing herbivores as an effective strategy for maintaining or bolstering reef resilience in a changing climate.

Investigating functional redundancy versus complementarity in Hawaiian herbivorous coral reef fishes.

Patterns of species resource use provide insight into the functional roles of species and thus their ecological significance within a community. The functional role of herbivorous fishes on coral reefs has been defined through a variety of methods, but from a grazing perspective, less is known about the species-specific preferences of herbivores on different groups of reef algae and the extent of dietary overlap across an herbivore community. Here, we quantified patterns of redundancy and complementarity in a highly diverse community of herbivores at a reef on Maui, Hawaii, USA. First, we tracked fish foraging behavior in situ to record bite rate and type of substrate bitten. Second, we examined gut contents of select herbivorous fishes to determine consumption at a finer scale. Finally, we placed foraging behavior in the context of resource availability to determine how fish selected substrate type. All species predominantly (73-100 %) foraged on turf algae, though there were differences among the types of macroalgae and other substrates bitten. Increased resolution via gut content analysis showed the composition of turf algae consumed by fishes differed across herbivore species. Consideration of foraging behavior by substrate availability revealed 50 % of herbivores selected for turf as opposed to other substrate types, but overall, there were variable foraging portfolios across all species. Through these three methods of investigation, we found higher complementarity among herbivorous fishes than would be revealed using a single metric. These results suggest differences across species in the herbivore "rain of bites" that graze and shape benthic community composition.

Decreased tourism during the COVID-19 pandemic positively affects reef fish in a high use marine protected area.

Humans alter ecosystems through both consumptive and non-consumptive effects. Consumptive effects occur through hunting, fishing and collecting, while non-consumptive effects occur due to the responses of wildlife to human presence. While marine conservation efforts have focused on reducing consumptive effects, managing human presence is also necessary to maintain and restore healthy ecosystems. Area closures and the tourism freeze related to the COVID-19 pandemic provided a unique natural experiment to measure the effects of decreased tourism on fish behavior in a high use no-take marine protected area (MPA) in Hawai`i. We found that when tourism shut down due to COVID restrictions in 2020, fish biomass increased and predatory species increased usage of shallow habitats, where tourists typically concentrate. When tourism resumed, fish biomass and habitat use returned to pre-pandemic levels. These displacement effects change fish community composition and biomass, which could affect key processes such as spawning, foraging and resting, and have knock-on effects that compromise ecosystem function and resilience. Managing non-consumptive uses, especially in heavily-visited MPAs, should be considered for sustainability of these ecosystems.

Displacement effects of heavy human use on coral reef predators within the Molokini Marine Life Conservation District.

The impact of marine ecotourism on reef predators is poorly understood and there is growing concern that overcrowding in Marine Protected Areas (MPAs) may disturb the species that these areas were established to protect. To improve our understanding of this issue, we used acoustic telemetry to examine the relationship between human activity at the Molokini Marine Life Conservation District (MLCD) and the habitat use of five reef-associated predators (Caranx melampygus, Caranx ignobilis, Triaenodon obesus, Carcharhinus amblyrhynchos, and Aprion virscens). During peak hours of human use, there was a negative relationship (R2=0.77, P<0.001) between the presence of bluefin trevally (Caranx melampygus) and vessels in subzone A. No other species showed strong evidence of this relationship. However, our results suggest that during this time, the natural ecosystem function that the reserve was established to protect may be compromised and overcrowding should be considered when managing MPAs.

Responses of Herbivorous Fishes and Benthos to 6 Years of Protection at the Kahekili Herbivore Fisheries Management Area, Maui.

In response to concerns about declining coral cover and recurring macroalgal blooms, in 2009 the State of Hawaii established the Kahekili Herbivore Fisheries Management Area (KHFMA). Within the KHFMA, herbivorous fishes and sea urchins are protected, but other fishing is allowed. As part of a multi-agency monitoring effort, we conducted surveys at KHFMA and comparison sites around Maui starting 19 months before closure, and over the six years since implementation of herbivore protection. Mean parrotfish and surgeonfish biomass both increased within the KHFMA (by 139% [95%QR (quantile range): 98-181%] and 28% [95%QR: 3-52%] respectively). Most of those gains were of small-to-medium sized species, whereas large-bodied species have not recovered, likely due to low levels of poaching on what are preferred fishery targets in Hawaii. Nevertheless, coincident with greater biomass of herbivores within the KHFMA, cover of crustose coralline algae (CCA) has increased from ~2% before closure to ~ 15% in 2015, and macroalgal cover has remained low throughout the monitoring period. Strong evidence that changes in the KHFMA were a consequence of herbivore management are that (i) there were no changes in biomass of unprotected fish families within the KHFMA; and that (ii) there were no similar changes in parrotfish or CCA at comparison sites around Maui. It is not yet clear how effective herbivore protection might eventually be for the KHFMA's ultimate goal of coral recovery. Coral cover declined over the first few years of surveys-from 39.6% (SE 1.4%) in 2008, to 32.9% (SE 0.8%) in 2012, with almost all of that loss occurring by 2010 (1 year after closure), i.e. before meaningful herbivore recovery had occurred. Coral cover subsequently stabilized and may have slightly increased from 2012 through early 2015. However, a region-wide bleaching event in 2015 had already led to some coral mortality by the time surveys were conducted in late 2015, at which time cover had dropped back to levels recorded in the KHFMA in 2012.

Effects of environmental salinity and 17alpha-methyltestosterone on growth and oxygen consumption in the tilapia, Oreochromis mossambicus.

Effects of environmental salinity and 17alpha-methyltestosterone (MT) on growth and oxygen consumption were examined in the tilapia, Oreochromis mossambicus. Yolk-sac fry were collected from brood stock in fresh water (FW). After yolk-sac absorption, they were assigned randomly to one of four groups: FW, MT treatment in FW, seawater (SW) and MT treatment in SW. All treatment groups were fed to satiation three times daily. The fish reared in SW (both control and MT-treated groups) grew significantly larger than either group in FW from day 43 throughout the experiment (195 days). The fish fed with MT added to their feed grew significantly larger than their respective controls from day 85 in FW and in SW until the end of the experiment. The routine metabolic rate (RMR) was determined monthly from month 2 (day 62) to month 5 (day 155). A significant negative correlation was seen between RMR and body mass in all treatment groups. Among fish of the same age, the SW-reared tilapia had significantly lower RMRs than the FW-reared fish. The MT-treated fish in SW showed significantly lower RMRs than the SW control group at months 3-5, whereas MT treatment in FW significantly increased the RMR at month 3. Comparison of regression lines between RMR and body mass indicates that MT treatment in FW caused a significant increase in oxygen consumption at a given mass of the fish, whereas MT treatment was without effect on RMR in SW-reared fish. These results clearly indicate that SW-rearing and MT treatment accelerate growth of tilapia, and that RMR decreases as fish size increased. It is also likely that the increased RMR and growth in MT-treated tilapia in FW may be due to the metabolic actions of MT, although the reason for the absence of MT treatment in SW is unclear.