Trade-offs between risks of predation and starvation in larvae make the shelf break an optimal spawning location for Atlantic bluefin tuna.

Atlantic bluefin tuna (ABT) (Thunnus thynnus) travel long distances to spawn in oligotrophic regions of the Gulf of Mexico (GoM) which suggests these regions offer some unique benefit to offspring survival. To better understand how larval survival varies within the GoM a spatially explicit, Lagrangian, individual-based model was developed that simulates dispersal and mortality of ABT early life stages within realistic predator and prey fields during the spawning periods from 1993 to 2012. The model estimates that starvation is the largest cumulative source of mortality associated with an early critical period. However, elevated predation on older larvae is identified as the main factor limiting survival to late postflexion. As a result, first-feeding larvae have higher survival on the shelf where food is abundant, whereas older larvae have higher survival in the open ocean with fewer predators, making the shelf break an optimal spawning area. The modeling framework developed in this study explicitly simulates both physical and biological factors that impact larval survival and hence could be used to support ecosystem based management efforts for ABT under current and future climate conditions.

Relationships between blooms of Karenia brevis and hypoxia across the West Florida Shelf.

Harmful algal blooms (HABs) caused by the dinoflagellate Karenia brevis on the West Florida Shelf have become a nearly annual occurrence causing widespread ecological and economic harm. Effects range from minor respiratory irritation and localized fish kills to large-scale and long-term events causing massive mortalities to marine organisms. Reports of hypoxia on the shelf have been infrequent; however, there have been some indications that some HABs have been associated with localized hypoxia. We examined oceanographic data from 2004 to 2019 across the West Florida Shelf to determine the frequency of hypoxia and to assess its association with known HABs. Hypoxia was present in 5 of the 16 years examined and was always found shoreward of the 50-meter bathymetry line. There were 2 clusters of recurrent hypoxia: midshelf off the Big Bend coast and near the southwest Florida coast. We identified 3 hypoxic events that were characterized by multiple conductivity, temperature, and depth (CTD) casts and occurred concurrently with extreme HABs in 2005, 2014, and 2018. These HAB-hypoxia events occurred when K. brevis blooms initiated in early summer months and persisted into the fall likely driven by increased biological oxygen demand from decaying algal biomass and reduced water column ventilation due to stratification. There were also four years, 2011, 2013, 2015, and 2017, with low dissolved oxygen located near the shelf break that were likely associated with upwelling of deeper Gulf of Mexico water onto the shelf. We had difficulty in assessing the spatiotemporal extent of these events due to limited data availability and potentially unobserved hypoxia due to the inconsistent difference between the bottom of the CTD cast and the seafloor. While we cannot unequivocally explain the association between extreme HABs and hypoxia on the West Florida Shelf, there is sufficient evidence to suggest a causal linkage between them.

Predicting ecosystem components in the Gulf of Mexico and their responses to climate variability with a dynamic Bayesian network model.

The Gulf of Mexico is an ecologically and economically important marine ecosystem that is affected by a variety of natural and anthropogenic pressures. These complex and interacting pressures, together with the dynamic environment of the Gulf, present challenges for the effective management of its resources. The recent adoption of Bayesian networks to ecology allows for the discovery and quantification of complex interactions from data after making only a few assumptions about observations of the system. In this study, we apply Bayesian network models, with different levels of structural complexity and a varying number of hidden variables to account for uncertainty when modeling ecosystem dynamics. From these models, we predict focal ecosystem components within the Gulf of Mexico. The predictive ability of the models varied with their structure. The model that performed best was parameterized through data-driven learning techniques and accounted for multiple ecosystem components' associations and their interactions with human and natural pressures over time. Then, we altered sea surface temperature in the best performing model to explore the response of different ecosystem components to increased temperature. The magnitude and even direction of predicted responses varied by ecosystem components due to heterogeneity in driving factors and their spatial overlap. Our findings suggest that due to varying components' sensitivity to drivers, changes in temperature will potentially lead to trade-offs in terms of population productivity. We were able to discover meaningful interactions between ecosystem components and their environment and show how sensitive these relationships are to climate perturbations, which increases our understanding of the potential future response of the system to increasing temperature. Our findings demonstrate that accounting for additional sources of variation, by incorporating multiple interactions and pressures in the model layout, has the potential for gaining deeper insights into the structure and dynamics of ecosystems.

Timing and locations of reef fish spawning off the southeastern United States.

Managed reef fish in the Atlantic Ocean of the southeastern United States (SEUS) support a multi-billion dollar industry. There is a broad interest in locating and protecting spawning fish from harvest, to enhance productivity and reduce the potential for overfishing. We assessed spatiotemporal cues for spawning for six species from four reef fish families, using data on individual spawning condition collected by over three decades of regional fishery-independent reef fish surveys, combined with a series of predictors derived from bathymetric features. We quantified the size of spawning areas used by reef fish across many years and identified several multispecies spawning locations. We quantitatively identified cues for peak spawning and generated predictive maps for Gray Triggerfish (Balistes capriscus), White Grunt (Haemulon plumierii), Red Snapper (Lutjanus campechanus), Vermilion Snapper (Rhomboplites aurorubens), Black Sea Bass (Centropristis striata), and Scamp (Mycteroperca phenax). For example, Red Snapper peak spawning was predicted in 24.7-29.0°C water prior to the new moon at locations with high curvature in the 24-30 m depth range off northeast Florida during June and July. External validation using scientific and fishery-dependent data collections strongly supported the predictive utility of our models. We identified locations where reconfiguration or expansion of existing marine protected areas would protect spawning reef fish. We recommend increased sampling off southern Florida (south of 27° N), during winter months, and in high-relief, high current habitats to improve our understanding of timing and location of reef fish spawning off the southeastern United States.

Location Isn't Everything: Timing of Spawning Aggregations Optimizes Larval Replenishment.

Many species of reef fishes form large spawning aggregations that are highly predictable in space and time. Prior research has suggested that aggregating fish derive fitness benefits not just from mating at high density but, also, from oceanographic features of the spatial locations where aggregations occur. Using a probabilistic biophysical model of larval dispersal coupled to a fine resolution hydrodynamic model of the Florida Straits, we develop a stochastic landscape of larval fitness. Tracking virtual larvae from release to settlement and incorporating changes in larval behavior through ontogeny, we found that larval success was sensitive to the timing of spawning. Indeed, propagules released during the observed spawning period had higher larval success rates than those released outside the observed spawning period. In contrast, larval success rates were relatively insensitive to the spatial position of the release site. In addition, minimum (rather than mean) larval survival was maximized during the observed spawning period, indicating a reproductive strategy that minimizes the probability of recruitment failure. Given this landscape of larval fitness, we take an inverse optimization approach to define a biological objective function that reflects a tradeoff between the mean and variance of larval success in a temporally variable environment. Using this objective function, we suggest that the length of the spawning period can provide insight into the tradeoff between reproductive risk and reward.

Evidence of climate-driven ecosystem reorganization in the Gulf of Mexico.

The Gulf of Mexico is one of the most ecologically and economically valuable marine ecosystems in the world and is affected by a variety of natural and anthropogenic phenomena including climate, hurricanes, coastal development, agricultural runoff, oil spills, and fishing. These complex and interacting stressors, together with the highly dynamic nature of this ecosystem, present challenges for the effective management of its resources. We analyze a compilation of over 100 indicators representing physical, biological, and economic aspects of the Gulf of Mexico and find that an ecosystem-wide reorganization occurred in the mid-1990s. Further analysis of fishery landings composition data indicates a major shift in the late 1970s coincident with the advent of US national fisheries management policy, as well as significant shifts in the mid-1960s and the mid-1990s. These latter shifts are aligned temporally with changes in a major climate mode in the Atlantic Ocean: the Atlantic Multidecadal Oscillation (AMO). We provide an explanation for how the AMO may drive physical changes in the Gulf of Mexico, thus altering higher-level ecosystem dynamics. The hypotheses presented here should provide focus for further targeted studies, particularly in regard to whether and how management should adjust to different climate regimes or states of nature. Our study highlights the challenges in understanding the effects of climatic drivers against a background of multiple anthropogenic pressures, particularly in a system where these forces interact in complex and nonlinear ways.

Spatial distribution and conservation of speckled hind and warsaw grouper in the Atlantic Ocean off the southeastern U.S.

There is broad interest in the development of efficient marine protected areas (MPAs) to reduce bycatch and end overfishing of speckled hind (Epinephelus drummondhayi) and warsaw grouper (Hyporthodus nigritus) in the Atlantic Ocean off the southeastern U.S. We assimilated decades of data from many fishery-dependent, fishery-independent, and anecdotal sources to describe the spatial distribution of these data limited stocks. A spatial classification model was developed to categorize depth-grids based on the distribution of speckled hind and warsaw grouper point observations and identified benthic habitats. Logistic regression analysis was used to develop a quantitative model to predict the spatial distribution of speckled hind and warsaw grouper as a function of depth, latitude, and habitat. Models, controlling for sampling gear effects, were selected based on AIC and 10-fold cross validation. The best-fitting model for warsaw grouper included latitude and depth to explain 10.8% of the variability in probability of detection, with a false prediction rate of 28-33%. The best-fitting model for speckled hind, per cross-validation, included latitude and depth to explain 36.8% of the variability in probability of detection, with a false prediction rate of 25-27%. The best-fitting speckled hind model, per AIC, also included habitat, but had false prediction rates up to 36%. Speckled hind and warsaw grouper habitats followed a shelf-edge hardbottom ridge from North Carolina to southeast Florida, with speckled hind more common to the north and warsaw grouper more common to the south. The proportion of habitat classifications and model-estimated stock contained within established and proposed MPAs was computed. Existing MPAs covered 10% of probable shelf-edge habitats for speckled hind and warsaw grouper, protecting 3-8% of speckled hind and 8% of warsaw grouper stocks. Proposed MPAs could add 24% more probable shelf-edge habitat, and protect an additional 14-29% of speckled hind and 20% of warsaw grouper stocks.

Adaptive significance of the formation of multi-species fish spawning aggregations near submerged capes.

BACKGROUND: Many fishes are known to spawn at distinct geomorphological features such as submerged capes or "promontories," and the widespread use of these sites for spawning must imply some evolutionary advantage. Spawning at these capes is thought to result in rapid offshore transport of eggs, thereby reducing predation levels and facilitating dispersal to areas of suitable habitat. METHODOLOGY/PRINCIPAL FINDINGS: To test this "off-reef transport" hypothesis, we use a hydrodynamic model and explore the effects of topography on currents at submerged capes where spawning occurs and at similar capes where spawning does not occur, along the Mesoamerican Barrier Reef. All capes modeled in this study produced eddy-shedding regimes, but specific eddy attributes differed between spawning and non-spawning sites. Eddies at spawning sites were significantly stronger than those at non-spawning sites, and upwelling and fronts were the products of the eddy formation process. Frontal zones, present particularly at the edges of eddies near the shelf, may serve to retain larvae and nutrients. Spawning site eddies were also more predictable in terms of diameter and longevity. Passive particles released at spawning and control sites were dispersed from the release site at similar rates, but particles from spawning sites were more highly aggregated in their distributions than those from control sites, and remained closer to shore at all times. CONCLUSIONS/SIGNIFICANCE: Our findings contradict previous hypotheses that cape spawning leads to high egg dispersion due to offshore transport, and that they are attractive for spawning due to high, variable currents. Rather, we show that current regimes at spawning sites are more predictable, concentrate the eggs, and keep larvae closer to shore. These attributes would confer evolutionary advantages by maintaining relatively similar recruitment patterns year after year.

Untangling natural seascape variation from marine reserve effects using a landscape approach.

Distinguishing management effects from the inherent variability in a system is a key consideration in assessing reserve efficacy. Here, we demonstrate how seascape heterogeneity, defined as the spatial configuration and composition of coral reef habitats, can mask our ability to discern reserve effects. We then test the application of a landscape approach, utilizing advances in benthic habitat mapping and GIS techniques, to quantify this heterogeneity and alleviate the confounding influence during reserve assessment. Seascape metrics were quantified at multiple spatial scales using a combination of spatial image analysis and in situ surveys at 87 patch reef sites in Glover's Reef Lagoon, Belize, within and outside a marine reserve enforced since 1998. Patch reef sites were then clustered into classes sharing similar seascape attributes using metrics that correlated significantly to observed variations in both fish and coral communities. When the efficacy of the marine reserve was assessed without including landscape attributes, no reserve effects were detected in the diversity and abundance of fish and coral communities, despite 10 years of management protection. However, grouping sites based on landscape attributes revealed significant reserve effects between site classes. Fish had higher total biomass (1.5x) and commercially important biomass (1.75x) inside the reserve and coral cover was 1.8 times greater inside the reserve, though direction and degree of response varied by seascape class. Our findings show that the application of a landscape classification approach vastly improves our ability to evaluate the efficacy of marine reserves by controlling for confounding effects of seascape heterogeneity and suggests that landscape heterogeneity should be considered in future reserve design.