Ocean warming has a greater effect than acidification on the early life history development and swimming performance of a large circumglobal pelagic fish.

Ocean warming and acidification are serious threats to marine life; however, their individual and combined effects on large pelagic and predatory fishes are poorly understood. We determined the effects of projected future temperature and carbon dioxide (CO2 ) levels on survival, growth, morphological development and swimming performance on the early life stages of a large circumglobal pelagic fish, the yellowtail kingfish Seriola lalandi. Eggs, larvae and juveniles were reared in cross-factored treatments of temperature (21 and 25°C) and pCO2 (500 and 985 µatm) from fertilisation to 25 days post hatching (dph). Temperature had the greatest effect on survival, growth and development. Survivorship was lower, but growth and morphological development were faster at 25°C, with surviving fish larger and more developed at 1, 11 and 21 dph. Elevated pCO2 affected size at 1 dph, but not at 11 or 21 dph, and did not affect survival or morphological development. Elevated temperature and pCO2 had opposing effects on swimming performance at 21 dph. Critical swimming speed (Ucrit ) was increased by elevated temperature but reduced by elevated pCO2 . Additionally, elevated temperature increased the proportion of individuals that responded to a startle stimulus, reduced latency to respond and increased maximum escape speed, potentially due to the more advanced developmental stage of juveniles at 25°C. By contrast, elevated pCO2 reduced the distance moved and average speed in response to a startle stimulus. Our results show that higher temperature is likely to be the primary driver of global change impacts on kingfish early life history; however, elevated pCO2 could affect critical aspects of swimming performance in this pelagic species. Our findings will help parameterise and structure fisheries population dynamics models and improve projections of impacts to large pelagic fishes under climate change scenarios to better inform adaptation and mitigation responses.

Elevated temperature and CO2 have positive effects on the growth and survival of larval Australasian snapper.

Rising water temperature and increased uptake of CO2 by the ocean are predicted to have widespread impacts on marine species. However, the effects are likely to vary, depending on a species' sensitivity and the geographical location of the population. Here, we investigated the potential effects of elevated temperature and pCO2 on larval growth and survival in a New Zealand population of the Australasian snapper, Chrysophyrs auratus. Eggs and larvae were reared in a fully cross-factored experiment (18 °C and 22 °C/pCO2 440 and 1040 µatm) to 16 days post hatch (dph). Morphologies at 1 dph and 16 dph were significantly affected by temperature, but not CO2. At 1dph, larvae at 22 °C were longer (7%) and had larger muscle depth at vent (14%), but had reduced yolk (65%) and oil globule size (16%). Reduced yolk reserves in recently hatched larvae suggests higher metabolic demands in warmer water. At 16 dph, larvae at elevated temperature were longer (12%) and muscle depth at vent was larger (64%). Conversely, survival was primarily affected by CO2 rather than temperature. Survivorship at 1 dph and 16 dph was 24% and 54% higher, respectively, under elevated CO2 compared with ambient conditions. Elevated temperature increased survival (24%) at 1 dph, but not at 16 dph. These results suggest that projected climate change scenarios may have an overall positive effect on early life history growth and survival in this population of C. auratus. This could benefit recruitment success, but needs to be weighed against negative effects of elevated CO2 on metabolic rates and swimming performance observed in other studies on the same population.

An uncertain future: Effects of ocean acidification and elevated temperature on a New Zealand snapper (Chrysophrys auratus) population.

Anthropogenic CO2 emissions are warming and acidifying Earth's oceans, which is likely to lead to a variety of effects on marine ecosystems. Fish populations will be vulnerable to this change, and there is now substantial evidence of the direct and indirect effects of climate change on fish. There is also a growing effort to conceptualise the effects of climate change on fish within population models. In the present study knowledge about the response of New Zealand snapper to warming and acidification was incorporated within a stock assessment model. Specifically, a previous tank experiment on larval snapper suggested both positive and negative effects, and otolith increment analysis on wild snapper indicated that growth may initially increase, followed by a potential decline as temperatures continue to warm. As a result of this uncertainty, sensitivity analysis was performed by varying average virgin recruitment (R0) by ±30%, adult growth by ±6%, but adjusting mean size at recruitment by +48% as we had better evidence for this increase. Overall adjustments to R0 had the biggest impact on the future yield (at a management target of 40% of an unfished population) of the Hauraki Gulf snapper fishery. The most negative scenario suggested a 29% decrease in fishery yield, while the most optimistic scenario suggested a 44% increase. While largely uncertain, these results provide some scope for predicting future impacts on the snapper fishery. Given that snapper is a species where the response to climate change has been specifically investigated, increasing uncertainty in a future where climate change and other stressors interact in complex and unpredictable ways is likely to be an important consideration for the management of nearly all fish populations.

Elevated CO2 affects anxiety but not a range of other behaviours in juvenile yellowtail kingfish.

Elevated seawater CO2 can cause a range of behavioural impairments in marine fishes. However, most studies to date have been conducted on small benthic species and very little is known about how higher oceanic CO2 levels could affect the behaviour of large pelagic species. Here, we tested the effects of elevated CO2, and where possible the interacting effects of high temperature, on a range of ecologically important behaviours (anxiety, routine activity, behavioural lateralization and visual acuity) in juvenile yellowtail kingfish, Seriola lalandi. Kingfish were reared from the egg stage to 25 days post-hatch in a full factorial design of ambient and elevated CO2 (~500 and ~1000 µatm pCO2) and temperature (21 °C and 25 °C). The effects of elevated CO2 were trait-specific with anxiety the only behaviour significantly affected. Juvenile S. lalandi reared at elevated CO2 spent more time in the dark zone during a standard black-white test, which is indicative of increased anxiety. Exposure to high temperature had no significant effect on any of the behaviours tested. Overall, our results suggest that juvenile S. lalandi are largely behaviourally tolerant to future ocean acidification and warming. Given the ecological and economic importance of large pelagic fish species more studies investigating the effect of future climate change are urgently needed.

Organ health and development in larval kingfish are unaffected by ocean acidification and warming.

Anthropogenic CO2 emissions are causing global ocean warming and ocean acidification. The early life stages of some marine fish are vulnerable to elevated ocean temperatures and CO2 concentrations, with lowered survival and growth rates most frequently documented. Underlying these effects, damage to different organs has been found as a response to elevated CO2 in larvae of several species of marine fish, yet the combined effects of acidification and warming on organ health are unknown. Yellowtail kingfish, Seriola lalandi, a circumglobal subtropical pelagic fish of high commercial and recreational value, were reared from fertilization under control (21 °C) and elevated (25 °C) temperature conditions fully crossed with control (500 µatm) and elevated (1,000 µatm) pCO2 conditions. Larvae were sampled at 11 days and 21 days post hatch for histological analysis of the eye, gills, gut, liver, pancreas, kidney and liver. Previous work found elevated temperature, but not elevated CO2, significantly reduced larval kingfish survival while increasing growth and developmental rate. The current histological analysis aimed to determine whether there were additional sublethal effects on organ condition and development and whether underlying organ damage could be responsible for the documented effects of temperature on survivorship. While damage to different organs was found in a number of larvae, these effects were not related to temperature and/or CO2 treatment. We conclude that kingfish larvae are generally vulnerable during organogenesis of the digestive system in their early development, but that this will not be exacerbated by near-future ocean warming and acidification.

Functional response of sport divers to lobsters with application to fisheries management.

Fishery managers must understand the dynamics of fishers and their prey to successfully predict the outcome of management actions. We measured the impact of a two-day exclusively recreational fishery on Caribbean spiny lobster in the Florida Keys, USA, over large spatial scales (>100 km) and multiple years and used a theoretical, predator-prey functional response approach to identify whether or not sport diver catch rates were density-independent (type I) or density-dependent (type II or III functional response), and if catch rates were saturated (i.e., reached an asymptote) at relatively high lobster densities. We then describe how this predator-prey framework can be applied to fisheries management for spiny lobster and other species. In the lower Keys, divers exhibited a type-I functional response, whereby they removed a constant and relatively high proportion of lobsters (0.74-0.84) across all pre-fishing-season lobster densities. Diver fishing effort increased in a linear manner with lobster prey densities, as would be expected with a type-I functional response, and was an order of magnitude lower in the upper Keys than lower Keys. There were numerous instances in the upper Keys where the density of lobsters actually increased from before to after the fishing season, suggesting some type of "spill-in effect" from surrounding diver-disturbed areas. With the exception of isolated reefs in the upper Keys, the proportion of lobsters removed by divers was density independent (type-I functional response) and never reached saturation at natural lobster densities. Thus, recreational divers have a relatively simple predatory response to spiny lobster, whereby catch rates increase linearly with lobster density such that catch is a reliable indicator of abundance. Although diver predation is extremely high (approximately 80%), diver predation pressure is not expected to increase proportionally with a decline in lobster density (i.e., a depensatory response), which could exacerbate local extinction. Furthermore, management actions that reduce diver effort should have a concomitant and desired reduction in catch. The recreational diver-lobster predator-prey construct in this study provides a useful predictive framework to apply to both recreational and commercial fisheries, and on which to build as management actions are implemented.

Do nursery habitats provide shelter from flow for juvenile fish?

Juvenile fish nurseries are an essential life stage requirement for the maintenance of many fish populations. With many inshore habitats globally in decline, optimising habitat management by increasing our understanding of the relationship between juvenile fish and nursery habitats may be a prudent approach. Previous research on post-settlement snapper (Chrysophrys auratus) has suggested that structure may provide a water flow refuge, allowing snapper to access high water flow sites that will also have a high flux of their pelagic prey. We investigated this hypothesis by describing how Artificial Seagrass Units (ASUs) modified water flow while also using a multi-camera set up to quantify snapper position in relation to this water flow environment. Horizontal water flow was reduced on the down-current side of ASUs, but only at the height of the seagrass canopy. While the highest abundance of snapper did occur down-current of the ASUs, many snapper also occupied other locations or were too high in the water column to receive any refuge from water flow. The proportion of snapper within the water column was potentially driven by strategy to access zooplankton prey, being higher on the up-current side of ASUs and on flood tides. It is possible that post-settlement snapper alternate position to provide opportunities for both feeding and flow refuging. An alternative explanation relates to an observed interaction between post-settlement snapper and a predator, which demonstrated that snapper can utilise habitat structure when threatened. The nature of this relationship, and its overall importance in determining the value of nursery habitats to post-settlement snapper remains an elusive next step.

Mechanisms explaining nursery habitat association: how do juvenile snapper (Chrysophrys auratus) benefit from their nursery habitat?

Nursery habitats provide elevated survival and growth to the organisms that associate with them, and as such are a crucial early life-stage component for many fishes and invertebrates. The exact mechanisms by which these benefits are afforded to associated organisms, however, are often unclear. Here we assessed potential explanations of the nursery function of structurally complex habitats for post-settlement snapper, Chrysophrys auratus, in New Zealand. Specifically, we deployed Artificial Seagrass Units (ASUs) and used a combination of video observation, netting and diet analysis of associated post-settlement snapper as well describing potential prey within the micro-habitats surrounding ASUs. We did not observe any predation attempts and few potential predators, suggesting that for snapper the nursery value of structurally complex habitats is not as a predation refuge. The diet of post-settlement snapper mostly consisted of calanoid and cyclopoid copepods, which were most commonly sampled from within the water column. Nearly all suspected feeding events were also observed within the water column. When considering the velocity of water flow at each ASU, plankton sampling revealed a greater availability of copepods with increasing current strength, while netting and video observation demonstrated that the abundance of snapper was highest at sites with intermediate water velocity. This study highlights that the interaction between water flow and food availability may represent an important trade-off between energy expenditure and food intake for post-settlement snapper. Structurally complex habitats may mediate this relationship, allowing snapper to access sites with higher food availability while reducing swimming costs. This mechanism may have broader relevance, potentially explaining the importance of estuarine nursery habitats for other species.