Justin R. Rizzari

Effects of Spearfishing on Reef Fish Populations in a Multi-Use Conservation Area

Although spearfishing is a popular method of capturing fish, its ecological effects on fish populations are poorly understood, which makes it difficult to assess the legitimacy and desirability of spearfishing in multi-use marine reserves. Recent management changes within the Great Barrier Reef Marine Park (GBRMP) fortuitously created a unique scenario by which to quantify the effects of spearfishing on fish populations. As such, we employed underwater visual surveys and a before-after-control-impact experimental design to investigate the effects of spearfishing on the density and size structure of target and non-target fishes in a multi-use conservation park zone (CPZ) within the GBRMP. Three years after spearfishing was first allowed in the CPZ, there was a 54% reduction in density and a 27% reduction in mean size of coral trout (Plectropomus spp.), the primary target species. These changes were attributed to spearfishing because benthic habitat characteristics and the density of non-target fishes were stable through time, and the density and mean size of coral trout in a nearby control zone (where spearfishing was prohibited) remained unchanged. We conclude that spearfishing, like other forms of fishing, can have rapid and substantial negative effects on target fish populations. Careful management of spearfishing is therefore needed to ensure that conservation obligations are achieved and that fishery resources are harvested sustainably. This is particularly important both for the GBRMP, due to its extraordinarily high conservation value and world heritage status, and for tropical island nations where people depend on spearfishing for food and income. To minimize the effects of spearfishing on target species and to enhance protection of functionally important fishes (herbivores), we recommend that fishery managers adjust output controls such as size- and catch-limits, rather than prohibit spearfishing altogether. This will preserve the cultural and social importance of spearfishing in coastal communities where it is practised.

Impact of conservation areas on trophic interactions between apex predators and herbivores on coral reefs

Apex predators are declining at alarming rates due to exploitation by humans, but we have yet to fully discern the impacts of apex predator loss on ecosystem function. In a management context, it is critically important to clarify the role apex predators play in structuring populations of lower trophic levels. Thus, we examined the top-down influence of reef sharks (an apex predator on coral reefs) and mesopredators on large-bodied herbivores. We measured the abundance, size structure, and biomass of apex predators, mesopredators, and herbivores across fished, no-take, and no-entry management zones in the Great Barrier Reef Marine Park, Australia. Shark abundance and mesopredator size and biomass were higher in no-entry zones than in fished and no-take zones, which indicates the viability of strictly enforced human exclusion areas as tools for the conservation of predator communities. Changes in predator populations due to protection in no-entry zones did not have a discernible influence on the density, size, or biomass of different functional groups of herbivorous fishes. The lack of a relationship between predators and herbivores suggests that top-down forces may not play a strong role in regulating large-bodied herbivorous coral reef fish populations. Given this inconsistency with traditional ecological theories of trophic cascades, trophic structures on coral reefs may need to be reassessed to enable the establishment of appropriate and effective management regimes.

Reassessing the trophic role of reef sharks as apex predators on coral reefs

Apex predators often have strong top-down effects on ecosystem components and are therefore a priority for conservation and management. Due to their large size and conspicuous predatory behaviour, reef sharks are typically assumed to be apex predators, but their functional role is yet to be confirmed. In this study, we used stomach contents and stable isotopes to estimate diet, trophic position and carbon sources for three common species of reef shark (Triaenodon obesus, Carcharhinus melanopterus and C. amblyrhynchos) from the Great Barrier Reef (Australia) and evaluated their assumed functional role as apex predators by qualitative and quantitative comparisons with other sharks and large predatory fishes. We found that reef sharks do not occupy the apex of coral reef food chains, but instead have functional roles similar to those of large predatory fishes such as snappers, emperors and groupers, which are typically regarded as high-level mesopredators. We hypothesise that a degree of functional redundancy exists within this guild of predators, potentially explaining why shark-induced trophic cascades are rare or subtle in coral reef ecosystems. We also found that reef sharks participate in multiple food webs (pelagic and benthic) and are sustained by multiple sources of primary production. We conclude that large conspicuous predators, be they elasmobranchs or any other taxon, should not axiomatically be regarded as apex predators without thorough analysis of their diet. In the case of reef sharks, our dietary analyses suggest they should be reassigned to an alternative trophic group such as high-level mesopredators. This change will facilitate improved understanding of how reef communities function and how removal of predators (e.g., via fishing) might affect ecosystem properties.

A test of trophic cascade theory: fish and benthic assemblages across a predator density gradient on coral reefs

Removal of predators is often hypothesized to alter community structure through trophic cascades. However, despite recent advances in our understanding of trophic cascades, evidence is often circumstantial on coral reefs because fishing pressure frequently co-varies with other anthropogenic effects, such as fishing for herbivorous fishes and changes in water quality due to pollution. Australia’s outer Great Barrier Reef (GBR) has experienced fishing-induced declines of apex predators and mesopredators, but pollution and targeting of herbivorous fishes are minimal. Here, we quantify fish and benthic assemblages across a fishing-induced predator density gradient on the outer GBR, including apex predators and mesopredators to herbivores and benthic assemblages, to test for evidence of trophic cascades and alternative hypotheses to trophic cascade theory. Using structural equation models, we found no cascading effects from apex predators to lower trophic levels: a loss of apex predators did not lead to higher levels of mesopredators, and this did not suppress mobile herbivores and drive algal proliferation. Likewise, we found no effects of mesopredators on lower trophic levels: a decline of mesopredators was not associated with higher abundances of algae-farming damselfishes and algae-dominated reefs. These findings indicate that top-down forces on coral reefs are weak, at least on the outer GBR. We conclude that predator-mediated trophic cascades are probably the exception rather than the rule in complex ecosystems such as the outer GBR.

Key aspects of the biology, fisheries and management of Coral grouper

Coral grouper (genus Plectropomus), or coral trout, are members of the grouper family (Epinephelidae) and are one of the largest and most conspicuous predatory fishes on Indo-Pacific coral reefs. They are highly-prized food fishes that are targeted by subsistence, artisanal, commercial and recreational fisheries throughout their geographic range. Plectropomus have broadly similar diets and habitat requirements to other tropical groupers, but typically have faster growth and higher natural mortality rates. Although these characteristics are expected to increase population turnover and reduce innate vulnerability to environmental and anthropogenic impacts relative to other groupers, many Plectropomus populations are in decline due to the combined effects of overfishing and habitat degradation. In many locations, stock depletion from uncontrolled fishing, particularly at spawning aggregation sites, has resulted in local fishery collapse. Therefore, improved management of wild populations is urgently required to ensure conservation and sustainable fisheries of Plectropomus. Where possible, a combination of no-take marine reserves, market-based management approaches, and allocation or resurrection of property rights systems are recommended to complement conventional fishery management actions that limit catch and effort. Additional investment in aquaculture propagation is also needed to reduce fishing pressure on wild stocks and support management initiatives. This global synthesis of information pertaining to the biology, fisheries and management of Plectropomus will assist in guiding future management actions that are attempting to address a range of stressors including fishing, reef habitat degradation, and the escalating effects of climate change.

Cooperative hunting and gregarious behaviour in the zebra lionfish, Dendrochirus zebra

Cooperative hunting is considered one of the most ubiquitous forms of cooperative behaviour in animals, and has been extensively studied in a range of taxa, including birds, mammals, fish and insects (Packer and Ruttan 1988). The zebra lionfish, Dendrochirus zebra (Cuvier 1829), feeds on small crustaceans and fishes and has been previously described as a solitary predator in which individuals hunt exclusively by themselves (Moyer and Zaiser 1981). Here, we report the occurrence of cooperative hunting in D. zebra from coral reefs around Lizard Island, northern Great Barrier Reef, Australia (14°40′S, 145°28′E). During September–December 2012, three different pairings (2–3 individuals) of D. zebra were observed cooperatively hunting in the field, therefore offering insight into a thus far unknown social behaviour in this species (Moyer and Zaiser 1981). Furthermore, hunting behaviour was filmed with video cameras (GoPro, Woodman Labs; see supplementary material) in a laboratory setting. As in the field (Fig. 1a), D. zebra pairs were observed hunting together on six different occasions in groups of 2–3 individuals. In each case, individuals would swim towards prey (cardinalfishes, Apogon doerderlini) undulating their dorsal spines, using their web-like fins as a visual-barrier, together herding prey into a confined area. Individuals would then take turns striking at their prey (Fig. 1b). On several occasions, one individual would proceed to produce a jet of water directed towards the prey (believed to confuse/distract prey increasing the chance of a successful predatory event; Albins and Lyons 2012). This mode of hunting has been documented for the invasive red lionfish, Pterois volitans (Albins and Lyons 2012), and is likely an idiosyncratic trait of lionfishes. Contrary to earlier suggestions that D. zebra is a social recluse (Moyer and Zaiser 1981), the current findings suggest that D. zebra exhibits gregarious behaviour, living together in groups of 3 (Fig. 1c). To our knowledge, this is the first documented case of intraspecific cooperative hunting and gregarious behaviour in lionfish. Recent findings (Lonnstedt and McCormick 2013) suggest that this hunting behavior is mirrored by P. volitans, which could have implications for hunting effectiveness of invasive lionfish in the Caribbean.

Observations of marine wildlife tourism effects on a non-focal species

A radio-acoustic positioning system was used to assess the effects of shark cage-diving operators (SCDO) on the fine-scale movements of a non-focal species, the smooth stingray Bathytoshia brevicaudata. The results revealed that the time spent in the array was individually variable, but generally increased when SCDO were present and that the presence of SCDO may have the capacity to elicit changes in the space use of B. brevicaudata. These results indicate that the effects of marine wildlife tourism may extend beyond the focal species of interest.

Decadal-scale response of detritivorous surgeonfishes (family Acanthuridae) to no-take marine reserve protection and changes in benthic habitat

No-take marine reserves (NTMR) are increasingly being implemented to mitigate the effects of fishing on coral reefs, yet determining the efficacy of NTMRs depends largely on partitioning the effects of fishing from the effect of benthic habitat. Species of coral-reef fishes typically decline in density when subjected to fishing or benthic disturbances, but this is not always the case. This study documents the long-term (8-31 years) response of six species of detritivorous surgeonfishes (family Acanthuridae) to NTMR protection and benthic habitat change at four islands (Apo, Sumilon, Mantigue, Selinog) in the central Philippines, each island with a NTMR and a monitored fished site. Despite being subject to moderate fishing pressure, these species did not increase in density with NTMR protection. However, density of these surgeonfishes had a strong negative relationship with cover of live hard coral and a strong positive relationship with cover of dead substratum (sand, rubble, hard dead substratum). These surgeonfishes typically feed over dead substrata and thus probably increase in density following large environmental disturbances that substantially reduce live hard coral cover. Here, we describe effects of environmental disturbance events (e.g., use of explosives, typhoons) that reduced live hard-coral cover and subsequent large increases (up to 25 fold) in surgeonfish densities, which then slowly (over 5-15 years) decreased in density as live hard coral recovered. Density of these functionally important surgeonfish species was influenced more by changes to benthic cover than by NTMR protection. Thus, we highlight the greater importance of bottom-up controls (i.e., benthic changes to food availability) than top-down control (i.e., fishing) on a functionally important group of coral-reef fishes.

Partitioning no-take marine reserve (NTMR) and benthic habitat effects on density of small and large-bodied tropical wrasses

No-take marine reserves (NTMRs) are increasingly implemented for fisheries management and biodiversity conservation. Yet, assessing NTMR effectiveness depends on partitioning the effects of NTMR protection and benthic habitat on protected species. Such partitioning is often difficult, since most studies lack well-designed sampling programs (i.e. Before-After-Control-Impact-Pair designs) spanning long-term time scales. Spanning 31 years, this study quantifies the effects of NTMR protection and changes to benthic habitat on the density of tropical wrasses (F. Labridae) at Sumilon and Apo Islands, Philippines. Five species of wrasse were studied: two species of large-bodied (40–50 cm TL) Hemigymnus that were vulnerable to fishing, and three species of small-bodied (10–25 cm TL) Thalassoma and Cirrhilabrus that were not targeted by fishing. NTMR protection had no measurable effect on wrasse density, irrespective of species or body size, over 20 (Sumilon) and 31 (Apo) years of protection. However, the density of wrasses was often affected strongly by benthic cover. Hemigymnus spp. had a positive association with hard coral cover, while Thalassoma spp. and Cirrhilabrus spp. had strong positive associations with cover of rubble and dead substratum. These associations were most apparent after environmental disturbances (typhoons, coral bleaching, crown of thorns starfish (COTS) outbreaks, use of explosives and drive nets) reduced live hard coral cover and increased cover of rubble, dead substratum and sand. Disturbances that reduced hard coral cover often reduced the density of Hemigymnus spp. and increased the density of Thalassoma spp. and Cirrhilabrus spp. rapidly (1–2 years). As hard coral recovered, density of Hemigymnus spp. often increased while density of Thalassoma spp. and Cirrhilabrus spp. often decreased, often on scales of 5–10 years. This study demonstrates that wrasse population density was influenced more by changes to benthic cover than by protection from fishing.

Decadal den fidelity of a highly mobile reef lobster

Palinurid (spiny) lobsters are a group of highly mobile decapod crustaceans (Childress and Jury 2006). An understanding of lobster behaviour and habitat use is valuable for fisheries management and conservation, especially since palinurids are among the world’s most sought-after seafood. The painted spiny lobster (Panulirus versicolor) exclusively inhabits coral reefs of the Indian and Pacific Oceans, including the Great Barrier Reef (GBR; Australia). Although some lobster species show fidelity to small reef areas, none are known to remain in the same place for periods up to a decade (Childress and Jury 2006). Here we report an observation of long-term (decadal) site fidelity of P. versicolor from Northwest Island (23°18’S, 151°43’E) in the GBR. On 3 May 2013, we captured a male P. versicolor (144 mm carapace length) that was initially captured on 28 December 2003 (114 mm carapace length). Identity was confirmed via elastomer implant and unique body markings (Fig. 1). Initial capture and final capture occurred at the same coral shelter or den (confirmed by GPS), although it was known occasionally to use other nearby dens (within 200 m). It shared a den with a small (two to three) harem of females, which changed through time. Despite being periodically harassed (i.e. captured and handled on at least five occasions between 2004 and 2006) and occasionally injured (i.e. limb loss due to autotomy), this individual maintained a long-term association with a particular den. Previous work has shown that individual spiny lobsters, including those of Jasus edwardsii and P. versicolor, are capable of moving 0.5–6 km (Kelly and MacDiarmid 2003; Frisch 2007). A high degree of site fidelity in marine species is often associated with a tendency towards over-exploitation and/or vulnerability to extinction (Cheung et al. 2007). Given the importance of the den for protection against potential predators and for providing mating opportunities, it is likely that protection of high quality dens (in marine reserves, for example) may provide excellent conservation outcomes for this species.