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Evaluating spatial management options for tiger shark (Galeocerdo cuvier) conservation in US Atlantic Waters
Abstract
There has been debate in the literature over the use and success of spatial management zones (i.e. marine protected areas and time/area closures) as policy tools for commercially exploited sharks. The tiger shark (Galeocerdo cuvier) is a highly migratory predator found worldwide in warm temperate and tropical seas, which is caught in multiple US fisheries. We used a spatially explicit modelling approach to investigate the impact of varying spatial management options in the Western North Atlantic Ocean on tiger shark biomass, catch, and distribution, and impacts to other species in the ecosystem. Results suggest that under current management scenarios, tiger shark biomass will increase over time. Model outputs indicate that protecting additional habitats will have relatively minimal impacts on tiger shark biomass, as would increasing or decreasing protections in areas not highly suitable for tiger sharks. However, increasing spatial management protections in highly suitable habitats is predicted to have a positive effect on their biomass. Results also predict possible spill-over effects from current spatial protections. Our results provide insights for evaluating differing management strategies on tiger shark abundance patterns and suggest that management zones may be an effective conservation tool for highly migratory species if highly suitable habitat is protected.
... It is interesting to note, however, that tiger sharks have consistently received greater attention in the North Atlantic when compared to the South region. For example, basic tiger shark biology data has been available for the North Atlantic since 1949 (Gudger, 1949), and several Northern hemisphere regions have made use of such a high available data volume to direct conservation actions (Driggers et al., 2008;Carlson et al., 2012;Hammerschlag et al., 2012;Graham et al., 2016;Sulikowski et al., 2016;Hammerschlag et al., 2017;Morgan et al., 2020). Underdeveloped and developing countries, however, have not matched these research efforts, due to a suite of issues, ranging from scarce or non-existent fishing regulations and fisheries statistics, lack of financial support, government funding cuts, and the still prohibitively high costs of several research methods and technologies in these regions, hindering shark fisheries management (Castello et al., 2009;Zerbe, 2013;Wade, 2016;MMA, 2014;Barreto et al., 2017;Reis-Filho & Leduc, 2017;Booth, Squires & Milner-Gulland, 2019;FAO, 2020;McManus & Neves, 2021). ...
... Reproduction data gaps significantly hinder conservation efforts, as they are paramount in proposing both adequate fisheries management and marine protected areas for specific ontogenetic phases (i.e., neonates, juveniles, or pregnant females), especially for nursery areas where neonates and juveniles are more susceptible to human pressure (Heupel et al., 2018;Morgan et al., 2020). We suggest urgent assessments on this topic, and that special attention be given to the southern limits of tiger shark distribution, as different fertility rates and gestational periods may occur due to the lower local temperatures in these regions (Yamaguchi, Taniuchi & Shimizu, 2000;Lombardi-Carlson et al., 2003;Taylor, Harry & Bennett, 2016) requiring differential management plans (Walker, 2005). ...
The tiger shark (Galeocerdo cuvier) has been relatively well assessed concerning biology and ecology aspects in both Atlantic and Pacific North America and in Caribbean waters. The amount of data in these regions has led to the species protection under capture quotas and with the creation of sanctuaries. The reality in developing countries, however, is the exact opposite, with scarce information on the species in the southern hemisphere, namely South American and African waters. In these regions, protection measures are insufficient, and studies on tiger shark biology and ecology are scarce, significantly hindering conservation and management efforts. Thus, the aim of this study was to compile scientific literature on the tiger shark in the South Atlantic and discuss the impact of these data (or lack thereof) distributed within a total of ten research categories for guiding management plans. In total, 41 scientific publications on different G. cuvier biology and ecology aspects were obtained. The most studied topics were Feeding Ecology (n = 12), followed by Human Interactions (n = 8), and Movements and Migration (n = 7). Northeastern Brazil (Southwest Atlantic) was the most researched area, probably due to the higher coastal abundance of tiger sharks in this area, alongside a high number of recorded attacks, justifying funding for studies in the region. No studies carried out in other South American or African countries were found. It is important to mention that even though some research topics are relatively well covered, a severe knowledge gap is noted for risk assessments and fisheries management, with a proposition for the implementation of sanctuaries noted. This is, however, particularly worrisome, as the South Atlantic is mostly unexplored in this regard for tiger sharks. It is also important to note how different the attention given to this species is in the North Atlantic when compared to the South region. Lastly, we highlight that the existence of sub-populations, the lack of migratory corridors geographically connecting distinct areas used by the species, and the lack of fisheries statistics on tiger shark landings, all increase the vulnerability of this species in the South Atlantic.
... Despite the propensity of this species to travel thousands of kilometers each year, readily cross jurisdictional boundaries (Lea et al., 2018), and be exposed to multiple fisheries (Graham et al., 2016) Bahamas. Recent modeling efforts suggest that the protection of highly suitable habitat, such as that in The Bahamas, will have a positive impact on the overall abundance of tiger sharks and potentially allow for spillover to adjacent areas (Morgan et al., 2020). ...
... Similarly, Immature female tiger sharks tagged in south Florida predominately remained along the continental shelf, while mature female tagged nearby in the western Bahamas often traveled thousands of kilometers (Hammerschlag et al., 2012) , using the protected waters of The Bahamas for 32% only of the time (Graham et al., 2016). Understanding these movements and habitat use criteria can be important for assessing the effectiveness of protective measures for regional tiger shark stocks (Morgan et al., 2020). ...
Abundances of large sharks are reported to have declined worldwide, and in response various levels of fisheries management and conservation efforts have been established. For example, Marine Protected Areas have been suggested as means to protect large expanses of ocean from fishing and other industrial activities (e.g., habitat destruction), and in 2011 The Commonwealth of The Bahamas established The Bahamas Shark Sanctuary. However, assessing the effectiveness of conservation efforts is challenging because consistent long‐term datasets of shark abundances are often lacking, especially throughout the Caribbean and The Bahamas. Here, we investigated the catch rates and demographics of tiger sharks Galeocerdo cuvier caught in a fishery‐independent survey near Bimini, The Bahamas, from 1984 to 2019 to assess relative abundance trends following the banning of longline fishing in 1993 and the subsequent establishment of the shark sanctuary. To contextualize the relative abundance trends near Bimini, we compared this to the relative abundance of tiger sharks in a fishery‐dependent survey from the Southeastern USA (SE USA), conducted from 1994 to 2019. Our data suggest that local abundance of tiger sharks has been stable near Bimini since the 1980s, including after the banning of longline fishing and the implementation of the shark sanctuary. In comparison, the abundance near the SE USA has slowly increased in the last decade, following potential declines in the decade preceding the USA Shark Management Plan. The results of this study provide some optimism that current conservation efforts in The Bahamas have been effective to maintain local tiger shark abundance within the protected area. Additionally, current fisheries management in the SE USA is allowing this species to recover within those waters. This article is protected by copyright. All rights reserved.
... Space use shifts can place species at risk even when successful management approaches have been implemented to conserve them (Davies et al., 2019;Santora et al., 2020). In the western North Atlantic, no specific conservation or management measures exist for tiger sharks, but vulnerability of this species to fisheries has been relatively low in recent decades due to overlap between their highly suitable habitat areas and conservation management zones that prohibit commercial longline fishing (Calich et al., 2018;Graham et al., 2016;Queiroz et al., 2019), which has likely contributed to their population recovery and growth (Morgan et al., 2020). Here, we found that spatial overlap between tracked tiger sharks and these protective zones varied from as much as 100% to as little as only 5% in response to ocean warming over the past decade. ...
... Indeed, previous modeling has predicted that tiger shark biomass in the western North Atlantic will be negatively affected under a scenario of decreased overlap between their highly suitable habitats and no-fishing zones (Morgan et al., 2020). ...
Given climate change threats to ecosystems, it is critical to understand the responses of species to warming. This is especially important in the case of apex predators since they exhibit relatively high extinction risk, and changes to their distribution could impact predator–prey interactions that can initiate trophic cascades. Here we used a combined analysis of animal tracking, remotely sensed environmental data, habitat modeling, and capture data to evaluate the effects of climate variability and change on the distributional range and migratory phenology of an ectothermic apex predator, the tiger shark (Galeocerdo cuvier). Tiger sharks satellite tracked in the western North Atlantic between 2010 and 2019 revealed significant annual variability in the geographic extent and timing of their migrations to northern latitudes from ocean warming. Specifically, tiger shark migrations have extended farther poleward and arrival times to northern latitudes have occurred earlier in the year during periods with anomalously high sea-surface temperatures. A complementary analysis of nearly 40 years of tiger shark captures in the region revealed decadal-scale changes in the distribution and timing of shark captures in parallel with long-term ocean warming. Specifically, areas of highest catch densities have progressively increased poleward and catches have occurred earlier in the year off the North American shelf. During periods of anomalously high sea-surface temperatures, movements of tracked sharks shifted beyond spatial management zones that had been affording them protection from commercial fishing and bycatch. Taken together, these study results have implications for fisheries management, human–wildlife conflict, and ecosystem functioning.
... Sin embargo, paradójicamente, las medidas que imponen prohibición total podrían no repercutir positivamente en las poblaciones que pretendían proteger, en la medida de que un gran porcentaje de capturas incidentales mueren antes de llegar al barco o después de ser liberados. Por ello, las políticas de gestión deberían de tomar, como complemento, un derrotero más "espacial" basado en la información científica disponible (ver capítulos 3, 4 y 7), protegiendo al menos, los segmentos de la población más vulnerables (Barker and Schluessel, 2005;Barnett et al., 2012;Letessier et al., 2017;Queiroz et al., 2019;Williamson et al., 2019;Morgan et al., 2020). ...
In the present work, key aspects of the biology and ecology of the shortfin mako were studied. Feeding habits, analysed in two ocean basins, indicated that pelagic fish and cephalopods were the main prey items. In the South Pacific Ocean, a marked sexual segregation was found, with females being more common in the SE region; this was also the area with a higher abundance of juveniles and of late-stage pregnant females. In the North Atlantic Ocean, large-scale horizontal movements (including trans-Atlantic migrations) were identified and diel vertical behaviour patterns described. Importantly, individuals that performed wider movements away from the tagging location were less at risk from surface longline fishing. Using tagging and recapture data that spanned a ten-year period, survival, dispersal, and fishing mortality rates for both mako and blue sharks were estimated. The presence of plastics and hooks was also observed for both species, in two studied ocean basins. Finally, bycatch rates for other internationally protected shark species that are commonly caught using surface longlines was estimated based on direct observations, which were several times higher than the official reported data. The results presented here are especially relevant for improving the management measures focused on pelagic sharks.
... For these species, local management is unlikely to mitigate the threats faced in the open ocean such as longline fishing. However, modelling approaches have demonstrated that No-Take Areas can have positive effects on numbers of large, mobile sharks, such as G. cuvier, if they encompass highly suitable habitats for these species 68,69 . Such benefits will depend upon the amount of time individual sharks spend within the boundaries and the life-history stages that experience this protection. ...
Quantifying the drivers of population size in reef sharks is critical for the development of appropriate conservation strategies. In north-west Australia, shark populations inhabit coral reefs that border growing centres of human population, industry, and tourism. However, we lack baseline data on reef sharks at large spatial scales (hundreds of km) that might enable managers to assess the status of shark populations in the face of future development in this region. Here, we examined the occurrence, abundance and behaviour of apex (Galeocerdo cuvier, Carcharhinus plumbeus) and reef (C. amblyrhynchos, C. melanopterus, Triaenodon obesus) sharks using > 1200 deployments of baited remote underwater stereo-video systems (stereo-BRUVs) across > 500 km of coastline. We found evidence for species-specific influences of habitat and fishing activities on the occurrence (probability of observation), abundance (MaxN) and behaviour of sharks (time of arrival to the stereo-BRUVs and likelihood of feeding). Although the presence of management zoning (No-take areas) made little difference to most species, C. amblyrhynchos were more common further from boat ramps (a proxy of recreational fishing pressure). Time of arrival for all species was also influenced by distance to boat ramp, although patterns varied among species. Our results demonstrate the capacity for behavioural metrics to complement existing measures of occurrence and abundance in assessing the potential impact of human activities on shark populations.
... However, despite the large decline of shark populations and the high extinction risk (Baum et al., 2003;Pacoureau et al., 2021), exploitation of the stocks of oceanic sharks has only recently been regulated in the North Atlantic (International Commission for the Conservation of Atlantic Tunas [ICCAT], 2020). Nonetheless, management measures of longline fisheries could be further improved by minimising shark mortality, for example, by implementing spatial management such as seasonal closures and marine protected areas (MPAs), enforcing adaptations to depth of hook deployment, and stronger selectivity of target species by fishing gear to reduce bycatch (Queiroz et al., 2016Morgan et al., 2020;Pacoureau et al., 2021). Knowing where and when sharks are more vulnerable to fishing, such as we show here for adult blue sharks, will be important for the implementation of novel dynamic approaches for assessing the status of threatened oceanic sharks . ...
In the pelagic environment diel vertical movements (DVM) are widespread across taxa, from zooplankton ascending from day-time depths into surface layers at night to avoid visual predators, to apex predators following prey movements to maximise foraging opportunities. The drivers of DVM in large predators such as pelagic sharks have only recently begun to be investigated in detail with the advent of sophisticated archival tags and high-resolution oceanographic datasets. In this study, we satellite tagged adult [>180 cm fork length, (FL)] blue sharks (Prionace glauca) in the North Atlantic Ocean to examine behavioural changes in response to the encountered environment, and therefore, to determine potential risks of capture using pelagic longline fisheries data. Although blue sharks recurrently use surface waters, cyclic diel behaviours were observed, with >95% of night-time spent above 250 m depth and variable day-time depth use. Hence, three different diel behaviours were identified during the tracking period: (i) regular normal DVM (nDVM) (dawn descent – dusk ascent, with over 90% of nighttime spent above 250 m, and between 5 and 50% of the day below this threshold); (ii) surface-oriented behaviour (occupation of surface waters both day and night), and (iii) deep depth-oriented nDVM [dawn descent – dusk ascent, with the majority (>50%) of daytime spent at depth]. Importantly, diel behaviours generally occurred in different ocean regions with nDVM frequently observed in high latitudes, associated with cold, highly productive waters (e.g., North Atlantic Current/Labrador Current convergence zone, West African upwelling area), while depth-oriented nDVM was observed in warm, oligotrophic areas. Thus, day-time occupation of shallow waters significantly increased with lower water temperature at depth (100 m), and with increasing concentration (and decreasing depth) of the chlorophyll a maximum. During nights of full moon blue sharks spent significantly more time in the depth range of longline hooks, while fishing effort and catches were also higher. We demonstrate that increased occupancy of surface layers driven by highly productive, cold waters and greater lunar illumination lead to higher capture risk. Understanding habitat-specific vulnerability to fishing in a commercially important pelagic shark species is essential for improving management and conservation measures.
Effective ocean management and conservation of highly migratory species depends on resolving overlap between animal movements and distributions and fishing effort. Yet, this information is lacking at a global scale. Here we show, using a big-data approach combining satellite-tracked movements of pelagic sharks and global fishing fleets, that 24% of the mean monthly space used by sharks falls under the footprint of pelagic longline fisheries. Space use hotspots of commercially valuable sharks and of internationally protected species had the highest overlap with longlines (up to 76% and 64%, respectively) and were also associated with significant increases in fishing effort. We conclude that pelagic sharks have limited spatial refuge from current levels of high-seas fishing effort. Results demonstrate an urgent need for conservation and management measures at high-seas shark hotspots and highlight the potential of simultaneous satellite surveillance of megafauna and fishers as a tool for near-real time, dynamic management.
Emerging conservation efforts for the world’s large predators may, if successful, restore natural predator–prey interactions. Marine reserves, where large predators tend to be relatively common, offer an experimental manipulation to investigate interactions between large-bodied marine predators and their prey. We hypothesized that southern stingrays—large, long-lived and highly interactive mesopredators—would invest in anti-predator behavior in marine reserves where predatory large sharks, the primary predator of stingrays, are more abundant. Specifically, we predicted southern stingrays in marine reserves would reduce the use of deep forereef habitats in the favor of shallow flats where the risk of shark encounters is lower. Baited remote underwater video was used to survey stingrays and reef sharks in flats and forereef habitats of two reserves and two fished sites in Belize. The interaction between “protection status” and “habitat” was the most important factor determining stingray presence. As predicted, southern stingrays spent more time interacting with baited remote underwater videos in the safer flats habitats, were more likely to have predator-inflicted damage inside reserves, and were less abundant in marine reserves but only in the forereef habitat. These results are consistent with a predation-sensitive habitat shift rather than southern stingray populations being reduced by direct predation from reef sharks. Our study provides evidence that roving predators can induce pronounced habitat shifts in prey that rely on crypsis and refuging, rather than active escape, in high-visibility, heterogeneous marine habitats. Given documented impacts of stingrays on benthic communities it is possible restoration of reef shark populations with reserves could induce reef ecosystem changes through behavior-mediated trophic cascades.
The timing and extent of international crossings by billfishes, tunas, and sharks in the Cuba-Mexico-United States (U.S.) triangle was investigated using electronic tagging data from eight species that resulted in >22,000 tracking days. Transnational movements of these highly mobile marine predators were pronounced with varying levels of bi- or tri-national population connectivity displayed by each species. Billfishes and tunas moved throughout the Gulf of Mexico and all species investigated (blue marlin, white marlin, Atlantic bluefin tuna, yellowfin tuna) frequently crossed international boundaries and entered the territorial waters of Cuba and/or Mexico. Certain sharks (tiger shark, scalloped hammerhead) displayed prolonged periods of residency in U.S. waters with more limited displacements, while whale sharks and to a lesser degree shortfin mako moved through multiple jurisdictions. The spatial extent of associated movements was generally associated with their differential use of coastal and open ocean pelagic ecosystems. Species with the majority of daily positions in oceanic waters off the continental shelf showed the greatest tendency for transnational movements and typically traveled farther from initial tagging locations. Several species converged on a common seasonal movement pattern between territorial waters of the U.S. (summer) and Mexico (winter).
The ability to predict animal movement based on environmental change is essential for understanding the dynamic nature of their spatial ecology, and in turn the effectiveness of conservation strategies. We used a large marine predator that displays partial migration (the tiger shark Galeocerdo cuvier) as a model to test the role of oceanic conditions in predicting the space-use of different size classes. By using generalized additive mixed models (GAMMs), we revealed that environmental variables (sea surface temperature, primary productivity, thermal fronts, and bathymetry) had much greater predictive power for the movements of large, migratory tiger sharks than for small, resident individuals. We also found that coverage of tiger shark movements within “shark sanctuaries” (protected areas specifically for sharks) in the northwest Atlantic could be increased from 12 to 52% through inclusion of Bermuda’s waters. However, as large tiger sharks are migratory, over 80% of potential longline fisheries interactions would still occur outside the boundaries of even the expanded protected areas. This emphasises that management of highly migratory species needs to be dynamic and account for changing interactions with fisheries over time, which in a changing climate may rely on predicting movements based on oceanic conditions to be effective.
The potential effectiveness of marine protected areas (MPAs) as a conservation tool for large sharks has been questioned due to the limited spatial extent of most MPAs in contrast to the complex life history and high mobility of many sharks. Here we evaluated the movement dynamics of a highly migratory apex predatory shark (tiger shark Galeocerdo cuvier) at the Galapagos Marine Reserve (GMR). Using data from satellite tracking passive acoustic telemetry, and stereo baited remote underwater video, we estimated residency, activity spaces, site fidelity, distributional abundances and migration patterns from the GMR and in relation to nesting beaches of green sea turtles (Chelonia mydas), a seasonally abundant and predictable prey source for large tiger sharks. Tiger sharks exhibited a high degree of philopatry, with 93% of the total satellite-tracked time across all individuals occurring within the GMR. Large sharks (> 200 cm TL) concentrated their movements in front of the two most important green sea turtle-nesting beaches in the GMR, visiting them on a daily basis during nocturnal hours. In contrast, small sharks (< 200 cm TL) rarely visited turtle-nesting areas and displayed diurnal presence at a third location where only immature sharks were found. Small and some large individuals remained in the three study areas even outside of the turtle-nesting season. Only two sharks were satellite-tracked outside of the GMR, and following long-distance migrations, both individuals returned to turtle-nesting beaches at the subsequent turtle-nesting season. The spatial patterns of residency and site fidelity of tiger sharks suggest that the presence of a predictable source of prey and suitable habitats might reduce the spatial extent of this large shark that is highly migratory in other parts of its range. This highly philopatric behaviour enhances the potential effectiveness of the GMR for their protection.
Knowledge of the diet and trophic ecology of apex predators is key for the implementation of effective ecosystem as well as species-based management initiatives. Using a combination of stomach content data and stable isotope analysis (δ¹⁵N and δ¹³C) the current study provides information on size-based and sex-specific variations in diet, trophic position (TP) and foraging habitat of tiger sharks (Galeocerdo cuvier) caught in the KwaZulu-Natal Sharks Board bather protection program. This study presents the longest time-series and most detailed analysis of stomach content data for G. cuvier worldwide. Prey identified from 628 non-empty stomachs revealed a size-based shift in diet. Reptiles, birds, mysticetes, and large shark species increased in dietary importance with G. cuvier size, concomitant with a decrease in smaller prey such as batoids and teleosts. Seasonal and decadal shifts in diet driven primarily by changes in the importance of elasmobranchs and mammal (cetacean) prey were recorded for medium sized (150–220 cm) G. cuvier. Both stomach content and stable isotope analysis indicated that G. cuvier is a generalist feeder at the population level. Size-based δ¹³C profiles indicated a movement to offshore foraging habitats by larger G. cuvier. Calculated TP varied by method ranging from 4.0 to 5.0 (TPSCA for stomach contents) and from 3.6 to 4.5 (TPscaled and TPadditive for δ¹⁵N). Large (> 220 cm) G. cuvier did not feed at discrete trophic levels, but rather throughout the food web. These data provide key information on the ecological role of G. cuvier to improve the accuracy of regional food web modelling. This will enable a better understanding of the ecological impacts related to changes in the abundance of this predator.
Relative abundance of many shark species in the Atlantic is assessed by compiling data from several independently conducted, but somewhat spatially limited surveys. Although these localized surveys annually sample the same populations, resulting trends in yearly indices often conflict with one another, thereby hindering interpretation of abundance patterns at broad spatial scales. We used delta-lognormal generalized linear models (GLMs) to generate indices of abundance for seven Atlantic coastal shark species from six fishery-independent surveys along the US east coast and Gulf of Mexico from 1975 to 2014. These indices were further analysed using dynamic factor analysis (DFA) to produce simplified, broad-scale common trends in relative abundance over the entire sampled distribution. Effects of drivers including the North Atlantic Oscillation index, the Atlantic Multidecadal Oscillation index, annually averaged sea surface temperature and species landings were evaluated within the DFA model. The two decadal oscillations and species landings were shown to affect shark distribution along south-east US coast. Estimated common trends of relative abundance for all large coastal shark species showed similar decreasing patterns into the early 1990s, periods of sustained low index values thereafter and recent indications of recovery. Small coastal shark species exhibited more regional variability in their estimated common trends, such that two common trends were required to adequately describe patterns in relative abundance throughout the Gulf of Mexico and Atlantic. Overall, all species’ (except the Gulf of Mexico blacknose shark) time series concluded with an increasing trend, suggestive of initial recovery from past exploitation.
As upper level predators, sharks serve an important role in marine ecosystems, but are often at risk from fisheries. Successful management of these species will require detailed information about their movements and distributions.
Using satellite telemetry, we investigated the long‐term horizontal movements and seasonal distributions of shortfin mako sharks Isurus oxyrinchus in the western North Atlantic Ocean.
Twenty‐six sharks (14 USA, 12 Mexico) were tracked for durations of 78–527 days. Ten sharks were tracked for >1 year. Sharks displayed region‐specific movements, with little distributional overlap between the Gulf of Mexico and Caribbean Sea, and the western North Atlantic. Sharks tagged off the USA ranged over a larger area, including shelf and pelagic habitats. Their core distribution was largely over the continental shelf and fluctuated seasonally, ranging from South Carolina, USA, in the winter to Nova Scotia, Canada, in the autumn, and appeared to follow seasonal productivity peaks while favouring warmer waters. Sharks tagged off Mexico displayed more restricted movements, largely confined to shelf habitats, with core activity centred year‐round on the eastern Campeche Bank, Mexico.
Sharks moved across the jurisdictional management boundaries of 17 nations, and the proportion of tracked sharks harvested (22%) was twice that obtained from previous fisheries‐dependent, conventional tagging studies.
Sharks also displayed considerable variability in movements, with seven sharks tagged off the USA making long‐distance, highly directional southern excursions into unproductive subtropical/tropical waters before returning north.
Policy implications . The large‐scale and region‐specific movements of shortfin mako sharks underscore the need for close cooperation amongst western North Atlantic nations and implementation of regionally and seasonally specific management strategies. The movement patterns also provide baseline information, which could be used in spatially explicit stock assessment models. Identification of high‐use areas by shortfin mako sharks provides focal areas for quantifying interactions with fisheries. The high harvest rate observed in our fisheries‐independent tracking study raises questions about the true rate of fisheries mortality experienced by shortfin mako sharks, calling for a cautionary interpretation of past stock assessments used to determine management policy for this highly migratory species of conservation concern.
The use of spatially explicit models (SEMs) in ecology has grown enormously in the past two decades. One major advancement has been that fine-scale details of landscapes, and of spatially dependent biological processes, such as dispersal and invasion, can now be simulated with great precision, due to improvements in computer technology. Many areas of modeling have shifted toward a focus on capturing these fine-scale details, to improve mechanistic understanding of ecosystems. However, spatially implicit models (SIMs) have played a dominant role in ecology, and arguments have been made that SIMs, which account for the effects of space without specifying spatial positions, have an advantage of being simpler and more broadly applicable, perhaps contributing more to understanding. We address this debate by comparing SEMs and SIMs in examples from the past few decades of modeling research. We argue that, although SIMs have been the dominant approach in the incorporation of space in theoretical ecology, SEMs have unique advantages for addressing pragmatic questions concerning species populations or communities in specific places, because local conditions, such as spatial heterogeneities, organism behaviors, and other contingencies, produce dynamics and patterns that usually cannot be incorporated into simpler SIMs. SEMs are also able to describe mechanisms at the local scale that can create amplifying positive feedbacks at that scale, creating emergent patterns at larger scales, and therefore are important to basic ecological theory. We review the use of SEMs at the level of populations, interacting populations, food webs, and ecosystems and argue that SEMs are not only essential in pragmatic issues, but must play a role in the understanding of causal relationships on landscapes.
The Gulf of Gabes is considered as one of the most productive areas of the southern Mediterranean Sea and it plays an important role in Tunisian economy. It is known to be an archetypal ecosystem in which the effects of fisheries are the most pronounced. Based on the stock assessment outcomes, it is as a highly exploited ecosystem. Thereupon, it becomes necessary to establish adequate measures to facilitate the recovery of the marine resources. The most important sets of management measures regard the establishment of Marine Protected Areas (MPAs). However, these management plans should be assessed beforehand to make sure of the relevance of the measure and its impact on marine resources. Modeling may significantly enhance our understanding of the likely impacts of fisheries management plans on groups that are very difficult to study and this approach gives insights at larger spatial scales. We used Ecospace to investigate the potential impacts of several spatial management plans on the ecosystem structure of the Gulf of Gabes. The Ecospace model is based on the existing Ecopath model elaborated by Hattab (2013). The simulation were carried over a 15-year period. The outcomes of the simulations, suggest that the implementation of MPAs in the Gulf of Gabes could be simultaneously beneficial for the ecosystem and fishing activities. However, the benefits are related to the characteristics of the MPA. The spatial simulations highlight that the location is crucial to the success of the MPA. Additionally, an increase in the size of a MPA can result in an increase in the spillover effect and, consequently, in catches in the neighborhood without harming ecosystem integrity. The configuration of the implemented MPA is of capital importance, a set of many small MPAs is more beneficial than fewer and larger MPAs, especially in terms of catches.
This study examines spillover effects resulting from US fishing regulations instituted to protect sea turtles. Sea turtles, along with US and foreign fisheries for swordfish co-occur on the high seas in the North and Central Pacific and that allows for “spillover effects.” When one fishery is required to curtail fishing activity to reduce incidental fishing mortality on sea turtle populations, the activity of other, unregulated fleets may change in ways that adversely affect the very species intended for protection. This study provides an empirical model that estimates these “spillover effects” on sea turtle bycatch resulting from production displacement between regulated US and less-regulated non-US fleets in the North and Central Pacific Ocean. The study demonstrates strong spillover effects, resulting in more sea turtle interaction due to increased foreign fleet activity when Hawaii swordfish production declines.
Study aim and location: Many populations of highly mobile marine fishes, including large sharks, are experiencing declines. The benefits of spatial management zones, such as marine protected areas (MPAs), for such animals are unclear. To help fill this knowledge gap, we examined core habitat use areas (CHUAs) for bull (Carcharhinus leucas), great hammerhead (Sphyrna mokarran) and tiger sharks (Galeocerdo cuvier) in relation to specific MPAs and exclusive economic zones (EEZs) in the western North Atlantic Ocean.
Methods: Bull, great hammerhead and tiger sharks (N = 86 total) were satellite tagged and tracked in southern Florida and the northern Bahamas between 2010 and 2013. Filtered and regularized positions from Argos locations of tag transmissions were used to generate CHUAs for these sharks. Overlaps of CHUAs with regional protected areas and exclusive economic management zones were quantified to determine the proportion of each tracked shark’s CHUA under spatial protection from exploitation.
Results: A total of 0%, 17.9% and 34.7% of the regional CHUAs for tracked bull, great hammerhead and tiger sharks, respectively, were fully protected from exploitation in the study area.
Main conclusions: Expansion of protected areas to include U.S. territorial waters would effectively protect 100% of the CHUAs for all tracked sharks in the study area. This finding is particularly significant for great hammerhead sharks, which are currently overfished, vulnerable to bycatch mortality and are the focus of strident regional conservation efforts. These findings also provide a
means to inform decision makers and marine conservation planning efforts as to the types of management actions available and potential efficacy of spatial protections for these marine predators.
The Northeast U.S. (NEUS) Continental Shelf Ecosystem is a dynamic environment. In order to evaluate the response of this ecosystem to numerous human-induced perturbations and to explore possible future scenarios, the Northeast Fisheries Science Center (NEFSC) instituted the Energy Modeling and Analysis eXercise (EMAX). The primary goal of EMAX was to establish an ecological network model (i.e., a nuanced energy budget) of the entire NEUS Ecosystem food web. The highly interdisciplinary EMAX work focused on four contemporary (1996-2000) subregions of the ecosystem; designated 36 network nodes (biomass state variables) across a broad range of the biological hierarchy; and incorporated a wide range of key rate
processes. The emphasis of EMAX was to explore the particular role of small pelagic fishes in the ecosystem, and various model configurations were constructed and psuedo-dynamic scenarios evaluated to explore how potential changes to this group can affect the rest of the food web. Preliminary results show that small pelagic fishes are clearly keystone species in the ecosystem. There are some differences across the four EMAX regions reflective of the local biology, but major patterns of network properties are similar over space. EMAX will continue to play a critical role in the further development of an ecosystem approach to fisheries (EAF) by acting as a catalogue of information and data; identifying major fluxes among biotic components of the ecosystem; serving as a basis for further analytical models; developing a way to evaluate biomass tradeoffs; and acting as a backdrop for a suite of other relevant management and research questions.
Significance
Shark populations are declining worldwide because of overexploitation by fisheries with unknown consequences for ecosystems. Although the harvest of oceanic sharks remains largely unregulated, knowing precisely where they interact with fishing vessels will better aid their conservation. We satellite track six species of shark and two entire longline fishing vessel fleets across the North Atlantic over multiple years. Sharks actively select and aggregate in space-use “hotspots” characterized by thermal fronts and high productivity. However, longline fishing vessels also target these habitats and efficiently track shark movements seasonally, leading to an 80% spatial overlap. Areas of highest overlap between sharks and fishing vessels show persistence between years, suggesting current hotspots are at risk, and arguing for introduction of international catch limits.
Advancing our knowledge of the reproductive biology and mating systems of freeranging sharks is inherently challenging. The large size and mobility of the specimens are just a few of the problems that make such studies complicated, and in some respects, impractical. The tiger shark Galeocerdo cuvier is a large, roving, apex predator found in many oceans throughout the world. Although their nomadic nature is primarily linked to large-scale migrations, these sharks can also display site fidelity. One site where this is known to occur is at Tiger Beach, Bahamas. Unique to the waters of this area is the consistent sighting of large females. While the sex-specific use of the area remains unknown, the shallow, warm environment could represent a critical habitat for reproductive events. To investigate the reproductive biology of tiger sharks at Tiger Beach, 65 individuals were opportunistically sampled between 2011 and 2014. Reproductive status of captured females (n = 59) was assessed with ultrasonography and by measuring circulating sex steroid hormones (progesterone, testosterone and estradiol). Our results indicate that Tiger Beach is a high-use site for female tiger sharks of mixed life stages. The results also suggest that Tiger Beach may function as a refuge habitat, allowing females to reach maturity free from male mating harassment, as well as functioning as a gestation ground where gravid females can benefit from year-round calm warm waters, which may reduce the gestation period and accelerate embryo development.
Long-distance movements of animals are an important driver of population spatial dynamics and determine the extent of overlap with area-focused human activities, such as fishing. Despite global concerns of declining shark populations, a major limitation in assessments of population trends or spatial management options is the lack of information on their long-term migratory behaviour. For a large marine predator, the tiger shark Galeocerdo cuvier, we show from individuals satellite-tracked for multiple years (up to 1101 days) that adult males undertake annually repeated, round-trip migrations of over 7,500 km in the northwest Atlantic. Notably, these migrations occurred between the highly disparate ecosystems of Caribbean coral reef regions in winter and high latitude oceanic areas in summer, with strong, repeated philopatry to specific overwintering insular habitat. Partial migration also occurred, with smaller, immature individuals displaying reduced migration propensity. Foraging may be a putative motivation for these oceanic migrations, with summer behaviour showing higher path tortuosity at the oceanic range extremes. The predictable migratory patterns and use of highly divergent ecosystems shown by male tiger sharks appear broadly similar to migrations seen in birds, reptiles and mammals, and highlight opportunities for dynamic spatial management and conservation measures of highly mobile sharks.
The value of no-take marine reserves as fisheries-management tools is controversial, particularly in high-poverty areas where human populations depend heavily on fish as a source of protein. Spillover, the net export of adult fish, is one mechanism by which no-take marine reserves may have a positive influence on adjacent fisheries. Spillover can contribute to poverty alleviation, although its effect is modulated by the number of fishermen and fishing intensity. In this study, we quantify the effects of a community-managed marine reserve in a high poverty area of Northern Mozambique. For this purpose, underwater visual censuses of reef fish were undertaken at three different times: 3 years before (2003), at the time of establishment (2006) and 6 years after the marine reserve establishment (2012). The survey locations were chosen inside, outside and on the border of the marine reserve. Benthic cover composition was quantified at the same sites in 2006 and 2012. After the reserve establishment , fish sizes were also estimated. Regression tree models show that the distance from the border and the time after reserve establishment were the variables with the strongest effect on fish abundance. The extent and direction of the spillover depends on trophic group and fish size. Poisson Generalized Linear Models show that, prior to the reserve establishment , the survey sites did not differ but, after 6 years, the abundance of all fish inside the reserve has increased and caused spillover of herbivorous fish. Spillover was detected 1km beyond the limit of the reserve for small herbivorous fishes. Six years after the establishment of a community-managed reserve, the fish assemblages have changed dramatically inside the reserve, and spillover is benefitting fish assemblages outside the reserve.
The degree of reef isolation is thought to limit the frequency of long-range dispersals in reef-associated sharks. Therefore, understanding how the behaviour and spatial ecology of a species differ across reef habitats is essential for developing sound conservation approaches. The present study examined the residency, movement and activity space of grey reef sharks (Carcharhinus amblyrhynchos) in the central Great Barrier Reef (GBR). An array of 56 acoustic receivers covering 17 semi-isolated coral reefs across 150 km was used to monitor shark movements. Forty C. amblyrhynchos were tagged with acoustic transmitters and monitored from 251-821 d. Most sharks were detected on a single reef; however, some individuals (4 females; 10 males) moved to up to five reefs. Residency index ranged from 0.02-1.0, with a mean ± SD of 0.78 ± 0.26. Mixed effect models showed that weekly and monthly residency was mainly influenced by shark size, with little or no effect of environmental parameters. Although C. amblyrhynchos were present year-round, juvenile sharks had lower residency to their tagging reef than adults. In addition, mature females were detected less between November and mid-February, which coincides with reported parturition in the central GBR. Long-term monitoring data revealed that C. amblyrhynchos exhibited high residency to their tagging reef, and therefore even in systems with semi-isolated reefs such as the GBR this species may benefit from spatial management approaches at the reef level. However, behavioural differences between sexes and life-stages of C. amblyrhynchos reported in this study suggest marine reserves may provide lower protection relative to remote and isolated coral reefs.
The net movement of individuals from marine reserves (also known as no-take marine protected areas) to the remaining fishing grounds is known as spillover and is frequently used to promote reserves to fishers on the grounds that it will benefit fisheries. Here we consider how mismanaged a fishery must be before spillover from a reserve is able to provide a net benefit for a fishery. For our model fishery, density of the species being harvested becomes higher in the reserve than in the fished area but the reduction in the density and yield of the fished area was such that the net effect of the closure was negative, except when the fishery was mismanaged. The extent to which effort had to exceed traditional management targets before reserves led to a spillover benefit varied with rates of growth and movement of the model species. In general, for well-managed fisheries, the loss of yield from the use of reserves was less for species with greater movement and slower growth. The spillover benefit became more pronounced with increasing mis-management of the stocks remaining available to the fishery. This model-based result is consistent with the literature of field-based research where a spillover benefit from reserves has only been detected when the fishery is highly depleted, often where traditional fisheries management controls are absent. We conclude that reserves in jurisdictions with well-managed fisheries are unlikely to provide a net spillover benefit.
Knowledge of the habitat use and migration patterns of large sharks is important for assessing the effectiveness of large predator Marine Protected Areas (MPAs), vulnerability to fisheries and environmental influences, and management of shark-human interactions. Here we compare movement, reef-fidelity, and ocean migration for tiger sharks, Galeocerdo cuvier, across the Coral Sea, with an emphasis on New Caledonia. Thirty-three tiger sharks (1.54 to 3.9 m total length) were tagged with passive acoustic transmitters and their localised movements monitored on receiver arrays in New Caledonia, the Chesterfield and Lord Howe Islands in the Coral Sea, and the east coast of Queensland, Australia. Satellite tags were also used to determine habitat use and movements among habitats across the Coral Sea. Sub-adults and one male adult tiger shark displayed year-round residency in the Chesterfields with two females tagged in the Chesterfields and detected on the Great Barrier Reef, Australia, after 591 and 842 days respectively. In coastal barrier reefs, tiger sharks were transient at acoustic arrays and each individual demonstrated a unique pattern of occurrence. From 2009 to 2013, fourteen sharks with satellite and acoustic tags undertook wide-ranging movements up to 1114 km across the Coral Sea with eight detected back on acoustic arrays up to 405 days after being tagged. Tiger sharks dove 1136 m and utilised three-dimensional activity spaces averaged at 2360 km(3). The Chesterfield Islands appear to be important habitat for sub-adults and adult male tiger sharks. Management strategies need to consider the wide-ranging movements of large (sub-adult and adult) male and female tiger sharks at the individual level, whereas fidelity to specific coastal reefs may be consistent across groups of individuals. Coastal barrier reef MPAs, however, only afford brief protection for large tiger sharks, therefore determining the importance of other oceanic Coral Sea reefs should be a priority for future research.
Marine ecosystem models are used to investigate marine protected area (MPA) benefits for coral reef ecosystems located in Raja Ampat, in the heart of the Coral Triangle. Field data from an integrated and diverse research project is used to develop a spatial ecosystem model using Ecopath, Ecosim, and Ecospace modelling software. The ecological and fisheries responses of a reef ecosystem to different levels of fishing effort restrictions inside MPAs are explored. The trade-offs of allowing some fisheries to operate inside the MPAs versus designating the MPAs as no-take zones are highlighted. The results show that rapid rebuilding of reef fish populations, especially the large charismatic species, requires no-take areas. Distinct trade-offs in spillover benefits are observed between partially fished and no-take MPAs.
An abnormally high shark attack rate verified off Recife could be related to migratory behavior of tiger sharks. This situation started after the construction of the Suape port to the south of Recife. A previous study suggested that attacking sharks could be following northward currents and that they were being attracted shoreward by approaching vessels. In this scenario, such northward movement pattern could imply a higher probability of sharks accessing the littoral area of Recife after leaving Suape. Pop-up satellite archival tags were deployed on five tiger sharks caught off Recife to assess their movement patterns off northeastern Brazil. All tags transmitted from northward latitudes after 7-74 days of freedom. The shorter, soak distance between deployment and pop-up locations ranged between 33-209 km and implied minimum average speeds of 0.02-0.98 km.h−1. Both pop-up locations and depth data suggest that tiger shark movements were conducted mostly over the continental shelf. The smaller sharks moved to deeper waters within 24 hours after releasing, but they assumed a shallower (< 50 m) vertical distribution for most of the monitoring period. While presenting the first data on tiger shark movements in the South Atlantic, this study also adds new information for the reasoning of the high shark attack rate verified in this region.
The Gulf of Maine (GOM) ecosystem has been observed to experience large changes over the last several decades, switching from a groundfish dominated system to a system dominated by crustacean species such as the American lobster (Homarus americanus) and crabs. The objective of this study is to develop a lobster ecosystem model and to evaluate the changes in the GOM ecosystem for the American lobster between 1980s and 1990s. A mass-balance ecosystem model was developed separately for the two time periods (1980s and 1990s) using Ecopath. The model includes 24 functional groups consisting of lobster, its key predator and prey species, and other ecologically important groups in the ecosystem such as zooplankton, phytoplankton, and detritus. Using the models developed we conducted a comparative analysis of trophic interactions and community structures of the GOM ecosystem for lobster in 1980s and 1990s. The balanced model suggests that the biomasses of shelled mollusk, crab, juvenile lobster, adult lobster, and Atlantic herring (Clupea harengus) were doubled, tripled or even more than tripled, but the groundfish biomass decreased substantially from 1980s to 1990s. The study shows that the trophic structures of the lobster ecosystem in the 1980s are different from those in the 1990s with a decrease in top predator biomass and an increase in the biomass of lower trophic level organisms. The ecosystem in 1990s was found to be more mature than in 1980s. This study suggests a regime switch in the GOM from high trophic groundfish dominated system to low trophic crustacean species dominated system during 1980s–1990s.
Little is known about the long-term movement patterns of most marine apex predators. A network of acoustic receivers was used to quantify the long-term movements of transmitter-equipped tiger sharks Galeocerdo cuvier Peron & Lesueur, 1822 in the Main Hawaiian Islands. Tiger sharks were wide-ranging, swam between islands and patrolled up to 109 km of contiguous coastline. Visits to specific acoustic receiver sites were typically brief (mean duration 3.3 min), unpredictable and interspersed by absences of weeks, months or years. This pattern may be an optimal foraging strategy for capturing risk-averse prey. Tiger sharks may have to move on soon after arriving in an area because the element of surprise is quickly lost and potential prey become wary and difficult to catch. Juvenile tiger sharks were significantly wider-ranging and less frequently detected than mature females. Juveniles may be avoiding predation by larger individuals, or exploring to find suitable home ranges. Tiger sharks may also switch movement patterns and foraging strategies to take advantage of different prey types, restricting their movements to exploit seasonally abundant and naive prey. Further empirical studies are required to directly link movement patterns with foraging.
1. There has been considerable debate over the past decade with respect to wildlife provisioning, especially resultant behavioural changes that may impact the ecological function of an apex predator. The controversy is exemplified by the shark diving industry, where major criticisms based on inference, anecdote and opinion stem from concerns of potential behaviourally mediated ecosystem effects because of ecotourism provisioning (aka‘chumming’ or feeding).
2. There is a general lack of empirical evidence to refute or support associated claims. The few studies that have investigated the behavioural impacts of shark provisioning ecotourism have generated conflicting conclusions, where the confidence in such results may suffer from a narrow spatial and temporal focus given the highly mobile nature of these predators. There is need for studies that examine the potential behavioural consequences of provisioning over ecologically relevant spatial and temporal scales.
3. To advance this debate, we conducted the first satellite telemetry study and movement analysis to explicitly examine the long-range migrations and habitat utilization of tiger sharks (Galeocerdo cuvier) originating in the Bahamas and Florida, two areas that differ significantly with regards to the presence/absence of provisioning ecotourism.
4. Satellite telemetry data rejected the behaviourally mediated effects of provisioning ecotourism at large spatial and temporal scales. In contrast, to the restricted activity space and movement that were hypothesized, geolocation data evidenced previously unknown long-distance migrations and habitat use for both tiger shark populations closely associated with areas of high biological productivity in the Gulf Stream and subtropical western Atlantic Ocean. We speculate that these areas are likely critically important for G. cuvier feeding forays and parturition.
5. We concluded that, in the light of potential conservation and public awareness benefits of ecotourism provisioning, this practice should not be dismissed out of hand by managers. Given the pressing need for improved understanding of the functional ecology of apex predators relative to human disturbance, empirical studies of different species sensitivities to disturbance should be used to guide best-practice ecotourism policies that maximize conservation goals.
Benefits for fisheries from marine protected areas (MPAs) are expected from recruitment of exported eggs and larvae as well as from spillover of adults to adjacent fishing grounds. Because the recruitment effect is difficult to detect, spillover is presently the only tangible potential fishery benefit of MPAs. Despite abundant evidence of spillover in the literature, this is the first study to quantify the number and biomass of individuals annually spilling over from an MPA and their contribution to the local fishery catches. Using a decade (1997-2007) of tag-recapture data for the lobster Palinurus elephas from the Columbretes Islands Marine Reserve (CIMR), we estimated annual emigration probabilities of 3.7% (female) and 6.7% (male), and quantified the resulting spillover to adjacent fished areas. We showed that during an 8 to 17 yr protection period, harvested spillover offset the loss of yield resulting from the reduction of fishing grounds set aside in the CIMR, producing a mean annual net benefit of 10% of the catch in weight. Although the number of lobsters spilling over annually did not quite make up for the loss of fishing grounds in the CIMR, it did in weight because the mean size of the lobsters emigrating from the reserve was larger than that of those outside. We propose that nomadic or home range movements of individuals near MPA boundaries, seasonal migrations and migrations forced by extraordinary meteorological events, and density-dependent movements facilitated spillover from the CIMR. Fishing effort concentration along its boundaries and high exploitation rates in the local fishery limited the spatial extent of spillover.
A simple model termed ECOPATH is pre- sented which estimates mean annual biomass, produc- tion, and consumption for components of an ecosystem. To use the model, the ecosystem must be partitioned into groups of similar species and provide for these species groups, estimates of production to biomass, diet, and food consumption. The ECOPATH model is applied to an ecosystem at French Frigate Shoals in the Northwest- ern Hawaiian Islands. Extensive field work provides both estimates of the input parameters as well as estimates of mean annual biomass and production. Biomass and pro- duction estimates for some of the species groups mod- eled are used to validate the estimates generated by the model.
Recovery of overexploited marine populations has been slow, and most remain below target biomass levels. A key question is
whether this is due to insufficient reductions in harvest rates or the erosion of population resilience. Using a global meta-analysis
of overfished stocks, we find that resilience of those stocks subjected to moderate levels of overfishing is enhanced, not
compromised, offering the possibility of swift recovery. However, prolonged intense overexploitation, especially for collapsed
stocks, not only delays rebuilding but also substantially increases the uncertainty in recovery times, despite predictable
influences of fishing and life history. Timely and decisive reductions in harvest rates could mitigate this uncertainty. Instead,
current harvest and low biomass levels render recovery improbable for the majority of the world’s depleted stocks.
During the reproductive season, sea turtles use a restricted area in the vicinity of their nesting beaches, making them vulnerable to predation. At Raine Island (Australia), the highest density green turtle Chelonia mydas rookery in the world, tiger sharks Galeocerdo cuvier have been observed to feed on green turtles, and it has been suggested that they may specialise on such air-breathing prey. However there is little information with which to examine this hypothesis. We compared the spatial and temporal components of movement behaviour of these two potentially interacting species in order to provide insight into the predator-prey relationship. Specifically, we tested the hypothesis that tiger shark movements are more concentrated at Raine Island during the green turtle nesting season than outside the turtle nesting season when turtles are not concentrated at Raine Island. Turtles showed area-restricted search behaviour around Raine Island for ∼3-4 months during the nesting period (November-February). This was followed by direct movement (transit) to putative foraging grounds mostly in the Torres Straight where they switched to area-restricted search mode again, and remained resident for the remainder of the deployment (53-304 days). In contrast, tiger sharks displayed high spatial and temporal variation in movement behaviour which was not closely linked to the movement behaviour of green turtles or recognised turtle foraging grounds. On average, tiger sharks were concentrated around Raine Island throughout the year. While information on diet is required to determine whether tiger sharks are turtle specialists our results support the hypothesis that they target this predictable and plentiful prey during turtle nesting season, but they might not focus on this less predictable food source outside the nesting season.
No-take marine protected areas (MPAs) are a commonly applied tool to reduce human fishing impacts on marine and coastal ecosystems. However, conservation outcomes of MPAs for mobile and long-lived predators such as sharks are highly variable. Here, we use empirical animal tracking data from 459 individual sharks and baited remote underwater video surveys undertaken in 36 countries to construct an empirically supported individual-based model that estimates the conservation effectiveness of MPAs for five species of coral reef-associated sharks (Triaenodon obesus, Carcharhinus melanopterus, Carcharhinus amblyrhynchos, Carcharhinus perezi, and Ginglymostoma cirratum). We demonstrate how species-specific individual movement traits can contribute to fishing mortality of sharks found within MPAs as they move outside to adjacent fishing grounds. We discovered that the world's officially recorded coral reef-based managed areas (with a median width of 9.4 km) would need to be enforced as strict no-take MPAs and up to 5 times larger to expect protection of the majority of individuals of the five investigated reef shark species. The magnitude of this effect depended on local abundances and fishing pressure, with MPAs required to be 1.6-2.6 times larger to protect the same number of Atlantic and Caribbean species, which occur at lower abundances than similar species in the western Pacific. Furthermore, our model was used to quantify partially substantial reductions (>50%) in fishing mortality resulting from small increases in MPA size, allowing us to bridge a critical gap between traditional conservation planning and fisheries management. Overall, our results highlight the challenge of relying on abundance data alone to ensure that estimates of shark conservation impacts of MPAs follow the precautionary approach.
Marine protected areas (MPAs) have grown exponentially, emerging as a widespread tool to conserve biodiversity and enhance fisheries production. Although numerous empirical studies and global syntheses have evaluated the effects of MPAs on community structure (e.g., biodiversity), no broad assessment concerning their capacity to influence ecological processes (e.g., species interactions) exists. Here, we present meta‐analyses that compare rates of predation and herbivory on a combined 32 species across 30 MPAs spanning 85° of latitude. Our analyses synthesize the fate of 15,225 field experiment assays, and demonstrate that MPAs greatly increased predation intensity on animals but not herbivory on macroalgae or seagrass. Predation risk, quantified as the odds of prey being eaten, was largely determined by predator abundance and biomass within reserves. At MPAs with the greatest predator accumulation, the odds of predation increased to nearly 49:1, as opposed to 1:1 at MPAs where predators actually declined. Surprisingly, we also found evidence that predation risk declined with increased sea‐surface temperature. Greater predation risk within MPAs was consistent with predator and prey population abundance estimates, where predators increased 4.4‐fold within MPAs, whereas prey decreased 2.2‐fold. For herbivory, the lack of change may have been driven by functional redundancy and the inability of reserves to increase herbivore abundance relative to fished zones in our sample. Overall, this work highlights the capacity of MPAs to restore a critical ecosystem function such as predation, which mediates energy flows and community assembly within natural systems. However, our review of the literature also uncovers relatively few studies that have quantified the effects of MPAs on ecosystem function, highlighting a key gap in our understanding of how protected areas may alter ecological processes and deliver ecosystem services. From a historical perspective, these findings suggest that modern levels of predation in the coastal oceans may currently only be a fraction of the baseline prior to human exploitation.
Arguments for the need to conserve aquatic predator (AP) populations often focus on the ecological and socioeconomic roles they play. Here, we summarize the diverse ecosystem functions and services connected to APs, including regulating food webs, cycling nutrients, engineering habitats, transmitting diseases/parasites, mediating ecological invasions, affecting climate, supporting fisheries, generating tourism, and providing bioinspiration. In some cases, human-driven declines and increases in AP populations have altered these ecosystem functions and services. We present a social ecological framework for supporting adaptive management decisions involving APs in response to social and environmental change. We also identify outstanding questions to guide future research on the ecological functions and ecosystem services of APs in a changing world.
Highly migratory species (e.g. sharks, tunas, turtles, cetaceans) present unique conservation
management challenges due to their wide-ranging movements. Consequently, the
extent to which management areas protect habitats for highly migratory species is often unknown. Within the southeast region of the USA’s exclusive economic zone, highly migratory sharks are target and/or bycatch species in pelagic and bottom longline fisheries. Here, we developed maximum entropy habitat suitability models for great hammerhead sharks Sphyrna mokarran, tiger sharks Galeocerdo cuvier, and bull sharks Carcharhinus leucas within the southeast region based on satellite tag (n = 96) and remotely sensed environmental data. Modeled highly suitable habitats were compared to longline gear management areas to determine what proportion of these habitats are protected from, and vulnerable to, longline fisheries. The percentages of highly suitable habitats overlapping with longline management areas varied by species and season (78% warm, 36% cool season for great hammerhead sharks; 48% warm, 79% cool for tiger sharks; and 2% warm, 100% cool for bull sharks). Highly suitable great hammerhead and tiger shark habitats were relatively well protected from pelagic longline fisheries yet vulnerable to bottom longline fisheries. Additionally, both species were vulnerable to pelagic and bottom longline fisheries off southwestern Florida; thus, extending gear restrictions to this area may benefit both species. Bull shark highly suitable habitats were only well protected from longline gear during the cool season. These results demonstrate how habitat suitability modeling can be used to help assess the efficacy of spatial management strategies and inform conservation plans for highly migratory species.
There is limited evidence on the rate at which the shark populations of coral reefs can rebound from over-exploitation, the baselines that might signify when recovery has occurred and the role of no-take Marine Protected Areas (MPA) in aiding this process. We surveyed shark assemblages at Ashmore Reef in Western Australia using baited remote underwater video stations in 2004 prior to enforcement of MPA status and then again in 2016 after eight years of strict enforcement. We found an increase in the relative mean abundance of Carcharhinus amblyrhynchos from 0.16 ± 0.06 individuals h−1 in 2004 to 0.74 ± 0.11 individuals h−1 in 2016, a change that was also accompanied by a shift in the assemblage of sharks to greater proportions of apex species (from 7.1% to 11.9%) and reef sharks (from 28.6% to 57.6%), and a decrease in the proportional abundance of lower trophic level species (from 64.3% to 30.5%). Abundances and trophic assemblage of sharks at Ashmore Reef in 2004 resembled those of the Scott Reefs, where targeted fishing for sharks still occurs, whereas in 2016, abundances and trophic structures had recovered to resemble those of the Rowley Shoals, a reef system that has been a strictly enforced MPA for over 25 years. The shift in abundance and community structure coincident with strict enforcement of the MPA at Ashmore Reef has occurred at a rate greater than predicted by demographic models, implying the action of compensatory processes in recovery. Our study shows that shark communities can recover rapidly after exploitation in a well-managed no-take MPA.
The redistribution of species has emerged as one of the most pervasive impacts of anthropogenic climate warming, and presents many societal challenges. Understanding how temperature regulates species distributions is particularly important for mobile marine fauna such as sharks given their seemingly rapid responses to warming, and the socio-political implications of human encounters with some dangerous species. The predictability of species distributions can potentially be improved by accounting for temperature’s influence on performance; an elusive relationship for most large animals. We combined multi-decadal catch data and bio-logging to show that coastal abundance and swimming performance of tiger sharks Galeocerdo cuvier are both highest at ~22°C, suggesting thermal constraints on performance may regulate this species’ distribution. Tiger sharks are responsible for a large proportion of shark bites on humans, and a focus of controversial control measures in several countries. The combination of distribution and performance data moves toward a mechanistic understanding of tiger shark’s thermal niche, and delivers a simple yet powerful indicator for predicting the location and timing of their occurrences throughout coastlines. For example, tiger sharks are mostly caught at Australia’s popular NSW beaches (i.e. near Sydney) in the warmest months, but our data suggest similar abundances will occur in winter and summer if annual sea surface temperatures increase by a further 1-2°C.
There is increasing concern for the conservation of sharks among scientists, environmental conservation advocates, and the interested public, but misunderstanding among policy non-specialists about which conservation and management policies are available, and which might work best for certain situations, persists. Here we present a review of fisheries management and conservation literature relating to sharks. Policies are broadly divided into target-based policies that aim for sustainable fisheries exploitation (e.g. fisheries quotas) and limit-based policies that aim to prevent all fisheries exploitation of entire taxa (e.g. marine reserves). A list of the pros and cons of each policy is included, as is a decision tree to aid in selection of the most appropriate policy. Our goal is that this paper will allow policy non-specialists, including scientists without policy training, environmental activists, and concerned citizens, to make informed decisions when advocating for shark conservation.
There is increasing concern about the conservation status of sharks. However, the presence of numerous different (and potentially mutually exclusive) policies complicates management implementation and public understanding of the process. Here we present the results of a survey that was distributed to members of the largest professional shark and ray research societies to assess member preferences for different conservation policies. Questions covered society member opinions towards available conservation and management policies, personal histories of getting involved in advocacy and management, and perceptions of the environmental conservation non-governmental organization (NGO) approach to shark conservation. Members of the shark and ray research community consider themselves to be knowledgeable about and actively involved in conservation and management policy. They are generally supportive of a variety of conservation policy tools that differ greatly in approach. Society members were generally less supportive of newer limit-based conservation policy tools, such as Shark Sanctuaries and shark fin bans, than of target-based fisheries management tools like fishing quotas that focus on species-specific sustainable exploitation. However, they provided few current examples of sustainable shark fisheries. Society members were generally supportive of environmental NGO efforts to conserve sharks, but raised concerns about some NGOs which are perceived as using incorrect information and focusing on the wrong problems. These results show that the ongoing debate in environmental policymaking circles between target-based natural resources management tools and limit-based conservation biology tools can also be found with respect to shark conservation and management. They also suggest that closer communication between the scientific and environmental NGO communities may be needed to recognize and reconcile differing values and objectives between these groups.
The ‘‘landscape of fear’’ model has been proposed as a unifying concept in ecology, describing, in part, how animals behave and move about in their environment. The basic model predicts that as an animal’s landscape changes from low to high risk of predation, prey species will alter their behavior to risk avoidance. However, studies investigating and evaluating the landscape of fear model across large spatial scales (tens to hundreds of thousands of square kilometers) in dynamic, open, aquatic systems involving apex predators and highly mobile prey are lacking. To address this knowledge gap, we investigated predator–prey relationships between tiger sharks (Galeocerdo cuvier) and loggerhead turtles (Caretta caretta) in the North Atlantic Ocean. This included the use of satellite tracking to examine shark and turtle distributions as well as their surfacing behaviors under varying levels of home range overlap. Our findings revealed patterns that deviated from our a priori predictions based on the landscape of fear model. Specifically, turtles did not alter their surfacing behaviors to risk avoidance when overlap in shark–turtle core home range was high. However, in areas of high overlap with turtles, sharks exhibited modified surfacing behaviors that may enhance predation opportunity. We suggest that turtles may be an important factor in determining shark distribution, whereas for turtles, other life history trade-offs may play a larger role in defining their habitat use. We propose that these findings are a result of both biotic and physically driven factors that independently or synergistically affect predator–prey interactions in this system. These results have implications for evolutionary biology, community ecology, and wildlife conservation. Further, given the difficulty in studying highly migratory marine species, our approach and conclusions may be applied to the study of other predator–prey systems.
This study used a network of acoustic receivers deployed around a no-take zone in Mangrove Bay, within the Ningaloo Reef Marine Park in Western Australia, to study residency and habitat preference of a small coastal shark, the nervous shark Carcharhinus cautus. Twelve C. cautus were tagged with acoustic tags and monitored for up to 579 days. Based on individuals detected within the receiver array for at least 2 months, C. cautus had small core (50% kernel utilization distribution, KUD) and home ranges (95% KUD) of 0·66 and 3·64 km(2) , respectively, and showed a strong habitat preference for mangroves, which are only found in the no-take zone. This resulted in C. cautus spending most of their detected time within the no-take zone boundaries (mean = 81·5%), showing that such a protected area could be beneficial to protect this species from extensive fishing pressure and local depletion, where required. Not all C. cautus remained within the acoustic array, however, suggesting that individual variations occur and that not all individuals would benefit from such protection. This study provides important information about the habitat, residency and movements of C. cautus that can be used for management and conservation. The strong affinity and residency of C. cautus within a mangrove-fringing coastline, emphasizes the importance of mangrove habitat to the species and suggests that such preferences can be used to design appropriate no-take zones for this species or others with similar habitat preferences.
© 2015 The Fisheries Society of the British Isles.
We examined various fishing management options to recover exploited marine resources and ecosystems of the Northern-Central Adriatic Sea. Dynamic simulations were based on a spatial ecological model previously calibrated with time series of data. Scenarios regarding spatial management were evaluated with the establishment of two marine protected areas, respectively, in the Pomo pit and the northern region. In addition, three temporal simulations of temporary closures and overall reduction of fishing effort of demersal and pelagic fleets (bottom, mid-water trawls and purse seines) were also considered. Simulations were run for 45 years (1975–2020), including the calibration period (1975–2002), and changes in biomass and catch of marine resources were analyzed. Our results confirm that current fishing management in the Adriatic Sea does not have clear beneficial impacts for the recovery of exploited resources, which will remain depleted in 2020 if “business as usual” continues. Simulations of alternative management suggest that both protected areas could be beneficial for fish population recovery predicting an increase in the biomass of commercial fish and predatory organisms. Simulations of temporary closures and overall reduction of fishing effort also show significant benefits for several commercial resources. We argue that both management measures may be effective tools to recover exploited ecosystems of the Northern-Central Adriatic Sea and halt the decline of marine resources.
Animals are often faced with complex movement decisions, particularly those that involve long-distance dispersal. Partial migrations, ubiquitous among all groups of vertebrates, are a form of long-distance movement that occurs when only some of the animals in a population migrate. The decision to migrate or to be a resident can be dependent on many factors, but these factors are rarely quantified in fishes, particularly top predators, even though partial migrations may have important implications for ecosystem dynamics and conservation. We utilized passive acoustic telemetry, with a Brownian bridge movement model and generalized additive mixed models, to explore the factors regulating partial migration in a large marine predator, the tiger shark, throughout the Hawaiian Islands. Although sharks tended to utilize a particular "core" island, they also demonstrated inter-island movements, particularly mature females that would swim from the northwestern Hawaiian Islands to the main Hawaiian Islands (MHI). Immigration to another island was a function of season, sea surface temperature (SST), and chlorophyll a concentration. Our results predict that 25% of mature females moved from remote French Frigate Shoals atoll to the MHI during late summer/early fall, potentially to give birth. Females with core home ranges within the MHI showed limited movements to the NWHI, and immigration to an island was better explained by SST and chlorophyll a concentration, suggesting a foraging function. Dispersal patterns in tiger sharks are complex but can be considered a mix of skipped-breeding partial migration by mature females and individual-based inter-island movements potentially linked to foraging. Therefore, sharks appear to use a conditional strategy based on fixed intrinsic and flexible extrinsic states. The application of Brownian bridge movement models to electronic presence/absence data provides a new technique for assessing the influence of habitat and environmental conditions on patterns of movement for fish populations.
Seasonal area closures of fisheries are primarily used to reduce fishing mortality on target species. In the absence of effort controls, fishing vessels displaced from a closed area will impact fish populations and the environment elsewhere. Based on the observed response of the North Sea beam trawl fleet to the closure of the ''cod box'' and an existing size-based model of the impacts of beam trawling, we predict the effects of seasonal area closures on benthic communities in the central North Sea. We suggest that repeated seasonal area closures would lead to a slightly more homogeneous distribution of annual trawling activity, although the distribution would remain patchy rather than random. The increased homogeneity, coupled with the displacement of trawling activity to previously unfished areas, is predicted to have slightly greater cumulative impacts on total benthic invertebrate production and lead to localized reductions in benthic biomass for several years. To en- sure the effective integration of fisheries and environmental management, the wider consequences of fishery management actions should be considered a priori. Thus, when seasonal closures increase the homogeneity of overall disturbance or lead to the re- distribution of trawling activity to environmentally sensitive or previously unfished areas, then effort reductions or permanent area closures should be considered as a manage- ment option. The latter would lead to a single but permanent redistribution of fishing disturbance, with lower cumulative impacts on benthic communities in the long run. 2003 International Council for the Exploration of the Sea. Published by Elsevier Science Ltd. All rights
Ecospace biomass-dynamics models for the central North Pacific predict strong space-time variation in abundances of various trophic groups in relation to nutrient-driven patterns in primary production and circulation-driven concentration of production in convergence areas. The model predicts simple patterns in ecosystem organization and abundances along productivity gradients. Predicted patterns are robust to alternative assumptions about how mobile organisms may alter dispersal behavior in relation to local fitness (per capita gain from net food intake minus predation mortality). Large marine protected areas (MPAs) would be needed to rebuild endangered populations and counter impacts of growing fishing effort. We expected that increases in dispersal rate in response to lower food availability and higher predation risk would reduce efficacy of MPAs as a management tool. Instead, simulations indicated that this negative effect may generally be eliminated or even reversed by positive effects of fitness-maximizing behaviors. Interannual variability in ocean circulation can further reduce the efficacy of MPAs; therefore, the dynamic nature of pelagic environments should also be considered in MPA design. Anomalies in ocean circulation may shift productive areas relative to MPA location, resulting in increased fishing mortality and (or) misinterpretation of catch statistics.
The widespread use of ecological network models (e.g., Ecopath, Econetwrk, and related energy budget models) has been laudable for several reasons, chief of which is providing an easy-to-use set of modeling tools that can present an ecosystem context for improved understanding and management of living marine resources (LMR). Yet the ease-of-use of these models has led to two challenges. First, the veritable explosion of the use and application of these network models has resulted in recognition that the content and use of such models has spanned a range of quality. Second, as these models and their application have become more widespread, they are increasingly being used in a LMR management context. Thus review panels and other evaluators of these models would benefit from a set of rigorous and standard criteria from which the basis for all network models and related applications for any given system (i.e., the initial, static energy budget) can be evaluated. To this end, as one suggestion for improving network models in general, here I propose a series of pre-balance (PREBAL) diagnostics. These PREBAL diagnostics can be done, now, in simple spreadsheets before any balancing or tuning is executed. Examples of these PREBAL diagnostics include biomasses, biomass ratios, vital rates, vital rate ratios, total production, and total removals (and slopes thereof) across the taxa and trophic levels in any given energy budget. I assert that there are some general ecological and fishery principles that can be used in conjunction with PREBAL diagnostics to identify issues of model structure and data quality before balancing and dynamic applications are executed. I humbly present this PREBAL information as a simple yet general approach that could be easily implemented, could be considered for further incorporation into these model packages, and as such would ultimately result in a straightforward way to evaluate (and perhaps identify areas for improving) initial conditions in food web modeling efforts.
From 1995 through 2006, 2577 bottom longline sets were conducted in the western North Atlantic Ocean from similar to 36 degrees N, 75 degrees W to 26 degrees N, 97 degrees W, an area that encompasses both the coastal waters off the southeastern USA (Atlantic) and the northern Gulf of Mexico (Gulf). Over the course of the study, 335 young-of-the-year (YOY) and 219 juvenile tiger sharks Galeocerdo cuvier were captured. In the Gulf, YOY catch per unit effort (CPUE) was highest from similar to 88 to 83 degrees W. In the Atlantic, the area of highest YOY CPUE occurred from 31 to 33 degrees N. Distribution of juvenile tiger sharks was more uniform than observed for YOY throughout the range of the study. Annual survival rates of YOY and Age 1+ tiger sharks were estimated to be 51 and 62 % in the Gulf and 39 and 27 % in the Atlantic. Total instantaneous mortality rates of YOY and Age 1+ tiger sharks were, respectively, estimated to be 0.67 and 0.47 in the Gulf and 0.93 and 1.32 in the Atlantic. In the Atlantic, age-specific natural mortality (M) estimates ranged from 0.12 to 0.50 and from 0.08 to 0.36 for YOY and juvenile tigers sharks, respectively. Estimates of age-specific Mwere more variable in the Gulf, with values ranging from 0.11 to 0.98 for YOY and from 0.08 to 0.57 for juveniles. In the absence of nursery areas, the production of numerous fast-growing offspring presumably helps the species survive in the face of high total mortality.