Michael J. Emslie

Rapid increase in fish numbers follows creation of world's largest marine reserve network

No-take marine reserves (NTMRs) are much advocated as a solution to managing marine ecosystems, protecting exploited species and restoring natural states of biodiversity [1,2]. Increasingly, it is becoming clear that effective marine conservation and management at ecosystem and regional scales requires extensive networks of NTMRs [1,2]. The worlds largest network of such reserves was established on Australias Great Barrier Reef (GBR) in 2004. Closing such a large area to all fishing has been socially and politically controversial, making it imperative that the effectiveness of this new reserve network be assessed. Here we report evidence, first, that the densities of the major target species of the GBR reef line fisheries were significantly higher in the new NTMRs, compared with fished sites, in just two years; and second, that the positive differences were consistent for multiple marine reserves over an unprecedented spatial scale (>1,000 km).

Maintenance of fish diversity on disturbed coral reefs

Habitat perturbations play a major role in shaping community structure; however, the elements of disturbance-related habitat change that affect diversity are not always apparent. This study examined the effects of habitat disturbances on species richness of coral reef fish assemblages using annual surveys of habitat and 210 fish species from 10 reefs on the Great Barrier Reef (GBR). Over a period of 11 years, major disturbances, including localised outbreaks of crown-of-thorns sea star (Acanthaster planci), severe storms or coral bleaching, resulted in coral decline of 46–96% in all the 10 reefs. Despite declines in coral cover, structural complexity of the reef framework was retained on five and species richness of coral reef fishes maintained on nine of the disturbed reefs. Extensive loss of coral resulted in localised declines of highly specialised coral-dependent species, but this loss of diversity was more than compensated for by increases in the number of species that feed on the epilithic algal matrix (EAM). A unimodal relationship between areal coral cover and species richness indicated species richness was greatest at approximately 20% coral cover declining by 3–4 species (6–8% of average richness) at higher and lower coral cover. Results revealed that declines in coral cover on reefs may have limited short-term impact on the diversity of coral reef fishes, though there may be fundamental changes in the community structure of fishes.

Spatial variation in the functional characteristics of herbivorous fish communities and the resilience of coral reefs

Many ecosystems face degradation unless factors that underpin their resilience can be effectively managed. In tropical reef ecosystems, grazing by herbivorous fishes can prevent coral-macroalgal phase shifts that commonly signal loss of resilience. However, knowledge of grazing characteristics that most promote resilience is typically experimental, localized, and sparse, which limits broad management applications. Applying sound ecological theory to broad-scale data may provide an alternative basis for ecosystem management. We explore the idea that resilience is positively related to the diversity within and among functional groups of organisms. Specifically, we infer the relative vulnerability of different subregions of the Great Barrier Reef (GBR) to phase shifts based on functional characteristics of the local herbivorous fish communities. Reef slopes on 92 reefs set in three zones of the continental shelf in eight latitudinal sectors of the GBR were surveyed on multiple occasions between 1995 and 2009. Spatial variation in fish community structure was high and driven primarily by shelf position. Measures of functional diversity, functional redundancy, and abundance were generally higher offshore and lower inshore. Two turbid inshore subregions were considered most vulnerable based on very low measures of herbivore function, and this was supported by the occurrence of phase shifts within one of three subregions. Eleven reefs that resisted phase shifts after major coral mortality included some with very low measures of herbivore function. The fact that phase shifts did not necessarily occur when large herbivores were scarce indicates that other environmental factors compensated to preserve resilience. Estimates of vulnerability based solely on herbivore function may thus prove conservative, but caution is appropriate, since compensatory factors are largely unknown and could be eroded unwittingly by anthropogenic stresses. Our data suggest that managing the threat of phase shifts in coral reef ecosystems successfully will require spatially explicit strategies that consider both the functional characteristics of local herbivore communities and environmental factors that may raise or lower resilience thresholds. A strong positive correlation between water clarity and the species richness and abundance of herbivorous fishes suggests that management of water quality is of generic importance to ensure the ecosystem services of this important group of herbivores.

Expectations and Outcomes of Reserve Network Performance following Re-zoning of the Great Barrier Reef Marine Park

Networks of no-take marine reserves (NTMRs) are widely advocated for preserving exploited fish stocks and for conserving biodiversity. We used underwater visual surveys of coral reef fish and benthic communities to quantify the short- to medium-term (5 to 30 years) ecological effects of the establishment of NTMRs within the Great Barrier Reef Marine Park (GBRMP). The density, mean length, and biomass of principal fishery species, coral trout (Plectropomus spp., Variola spp.), were consistently greater in NTMRs than on fished reefs over both the short and medium term. However, there were no clear or consistent differences in the structure of fish or benthic assemblages, non-target fish density, fish species richness, or coral cover between NTMR and fished reefs. There was no indication that the displacement and concentration of fishing effort reduced coral trout populations on fished reefs. A severe tropical cyclone impacted many survey reefs during the study, causing similar declines in coral cover and fish density on both NTMR and fished reefs. However, coral trout biomass declined only on fished reefs after the cyclone. The GBRMP is performing as expected in terms of the protection of fished stocks and biodiversity for a developed country in which fishing is not excessive and targets a narrow range of species. NTMRs cannot protect coral reefs directly from acute regional-scale disturbance but, after a strong tropical cyclone, impacted NTMR reefs supported higher biomass of key fishery-targeted species and so should provide valuable sources of larvae to enhance population recovery and long-term persistence.

Marine protected areas increase resilience among coral reef communities

With marine biodiversity declining globally at accelerating rates, maximising the effectiveness of conservation has become a key goal for local, national and international regulators. Marine protected areas (MPAs) have been widely advocated for conserving and managing marine biodiversity yet, despite extensive research, their benefits for conserving non-target species and wider ecosystem functions remain unclear. Here, we demonstrate that MPAs can increase the resilience of coral reef communities to natural disturbances, including coral bleaching, coral diseases, Acanthaster planci outbreaks and storms. Using a 20-year time series from Australias Great Barrier Reef, we show that within MPAs, (1) reef community composition was 21-38% more stable; (2) the magnitude of disturbance impacts was 30% lower and (3) subsequent recovery was 20% faster that in adjacent unprotected habitats. Our results demonstrate that MPAs can increase the resilience of marine communities to natural disturbance possibly through herbivory, trophic cascades and portfolio effects.

Great Barrier Reef butterflyfish community structure: the role of shelf position and benthic community type

The extent to which fish communities are structured by spatial variability in coral reef habitats versus stochastic processes (such as larval supply) is very important in predicting responses to sustained and ongoing habitat degradation. In this study, butterflyfish and benthic communities were surveyed annually over 15 years on 47 reefs (spanning 12° of latitude) of the Great Barrier Reef (GBR). Spatial autocorrelation in the structure of butterflyfish communities versus key differences in reef habitats was investigated to assess the extent to which the structure of these fish communities is influenced by habitat conditions. Benthic communities on each of the 47 reefs were broadly categorised as either: 1. Poritidae/Alcyoniidae, 2. mixed taxa, 3. soft coral or 4. Acropora-dominated habitats. These habitat types most reflected increases in water clarity and wave exposure, moving across the GBR shelf from coastal to outer-shelf environments. In turn, each habitat type also supported very distinct butterflyfish communities. Hard coral feeders were always the dominant butterflyfish species in each community type. However, the numerically dominant species changed according to habitat type, representing spatial replacement of species across the shelf. This study reveals clear and consistent differences in the structure of fish communities among reefs associated with marked differences in habitat structure.

Retention of Habitat Complexity Minimizes Disassembly of Reef Fish Communities following Disturbance: A Large-Scale Natural Experiment

High biodiversity ecosystems are commonly associated with complex habitats. Coral reefs are highly diverse ecosystems, but are under increasing pressure from numerous stressors, many of which reduce live coral cover and habitat complexity with concomitant effects on other organisms such as reef fishes. While previous studies have highlighted the importance of habitat complexity in structuring reef fish communities, they employed gradient or meta-analyses which lacked a controlled experimental design over broad spatial scales to explicitly separate the influence of live coral cover from overall habitat complexity. Here a natural experiment using a long term (20 year), spatially extensive (∼115,000 kms2) dataset from the Great Barrier Reef revealed the fundamental importance of overall habitat complexity for reef fishes. Reductions of both live coral cover and habitat complexity had substantial impacts on fish communities compared to relatively minor impacts after major reductions in coral cover but not habitat complexity. Where habitat complexity was substantially reduced, species abundances broadly declined and a far greater number of fish species were locally extirpated, including economically important fishes. This resulted in decreased species richness and a loss of diversity within functional groups. Our results suggest that the retention of habitat complexity following disturbances can ameliorate the impacts of coral declines on reef fishes, so preserving their capacity to perform important functional roles essential to reef resilience. These results add to a growing body of evidence about the importance of habitat complexity for reef fishes, and represent the first large-scale examination of this question on the Great Barrier Reef.

Effects of different disturbance types on butterflyfish communities of Australia's Great Barrier Reef

The effects of disturbances on coral reef fishes have been extensively documented but most studies have relied on opportunistic sampling following single events. Few studies have the spatial and temporal extent to directly compare the effects of multiple disturbances over a large geographic scale. Here, benthic communities and butterflyfishes on 47 reefs of the Great Barrier Reef were surveyed annually to examine their responses to physical disturbances (cyclones and storms) and/or biological disturbances (bleaching, outbreaks of crown-of-thorns starfish and white syndrome disease). The effects on benthic and butterflyfish communities varied among reefs depending on the structure and geographical setting of each community, on the size and type of disturbance, and on the disturbance history of that reef. There was considerable variability in the response of butterflyfishes to different disturbances: physical disturbances (occurring with or without biological disturbances) produced substantial declines in abundance, whilst biological disturbances occurring on their own did not. Butterflyfishes with the narrowest feeding preferences, such as obligate corallivores, were always the species most affected. The response of generalist feeders varied with the extent of damage. Wholesale changes to the butterflyfish community were only recorded where structural complexity of reefs was drastically reduced. The observed effects of disturbances on butterflyfishes coupled with predictions of increased frequency and intensity of disturbances sound a dire warning for the future of butterflyfish communities in particular and reef fish communities in general.

Patterns of embryo mortality in a demersally spawning coral reef fish and the role of predatory fishes

The biological significance of embryo mortality in demersally spawning coral reef fishes is poorly understood. Here we describe patterns of variation in embryo mortality in Pomacentrus amboinensis (Pomacentridae) at Lizard Island (Great Barrier Reef). The aim was to determine whether numbers of embryos hatched substantially differed from egg production, and if so, identify whether predatory fishes were a source of embryo mortality. Spawning success (number of eggs laid), embryo mortality (proportion of embryos that died prior to hatching) and number of embryos hatching were estimated from daily maps of clutches laid on artificial surfaces (PVC tiles) defended by nesting males. Patterns of variation in eggs laid, embryo mortality and numbers of embryos hatched were examined at three spatial scales: (1) among widely-spaced locations around the island; (2) between adjacent reef slope and patch reef habitats occupied by P. amboinensis at a single location; and (3) among different males within these two habitats. The embryo mortality was extremely high, with a mean of 25.9pm ± 6.2% (S.E.) for 4 locations examined in 1994 and a mean of 69.2pm ± 2.9% for two habitats surveyed in 1995. There were no significant differences in embryo mortality among locations or habitats in either year. This meant that spatial patterns in the number of embryos hatching reflected differences in the number of eggs laid on tiles. Embryo mortality was extremely variable on the scale of individual territories, with embryo mortality commonly ranging from 0% to 100%. Much of the mortality could be attributed to diurnal predatory fishes, especially the wrasse Thalassoma lunare. However, variation in predator densities did not explain spatial patterns in embryo mortality rates. Both solitary and group predatory behaviour was observed, with groups often causing 100% embryo mortality. The level of embryo mortality observed suggests that predation prior to hatching may have a substantial effect on the reproductive output of populations of this demersal-nesting fish.

Post-disturbance stability of fish assemblages measured at coarse taxonomic resolution masks change at finer scales

Quantifying changes to coral reef fish assemblages in the wake of cyclonic disturbances is challenging due to spatial variability of damage inherent in such events. Often, fish abundance appears stable at one spatial scale (e.g. reef-wide), but exhibits substantial change at finer scales (e.g. site-specific decline or increase). Taxonomic resolution also plays a role; overall stability at coarse taxonomic levels (e.g. family) may mask species-level turnover. Here we document changes to reef fish communities after severe Tropical Cyclone Ita crossed Lizard Island, Great Barrier Reef. Coral and reef fish surveys were conducted concurrently before and after the cyclone at four levels of exposure to the prevailing weather. Coral cover declined across all exposures except sheltered sites, with the largest decline at exposed sites. There was no significant overall reduction in the total density, biomass and species richness of reef fishes between 2011 and 2015, but individual fish taxa (families and species) changed in complex and unpredictable ways. For example, more families increased in density and biomass than decreased following Cyclone Ita, particularly at exposed sites whilst more fish families declined at lagoon sites even though coral cover did not decline. All sites lost biomass of several damselfish species, and at most sites there was an increase in macroinvertivores and grazers. Overall, these results suggest that the degree of change measured at coarse taxonomic levels masked high species-level turnover, although other potential explanations include that there was no impact of the storm, fish assemblages were impacted but underwent rapid recovery or that there is a time lag before the full impacts become apparent. This study confirms that in high-complexity, high diversity ecosystems such as coral reefs, species level analyses are essential to adequately capture the consequences of disturbance events.