Intraspecific variation in muscle growth of two distinct populations of Port Jackson sharks under projected end-of-century temperatures.

Although pervasive, the effects of climate change vary regionally, possibly resulting in differential behavioral, physiological, and/or phenotypic responses among populations within broadly distributed species. Juvenile Port Jackson sharks (Heterodontus portusjacksoni) from eastern and southern Australia were reared at their current (17.6 °C Adelaide, South Australia [SA]; 20.6 °C Jervis Bay, New South Wales [NSW]) or projected end-of-century (EOC) temperatures (20.6 °C Adelaide, SA; 23.6 °C Jervis Bay, NSW) and assessed for morphological features of skeletal muscle tissue. Nearly all skeletal muscle properties including cellularity, fiber size, myonuclear domain, and satellite cell density did not differ between locations and thermal regimes. However, capillary density was significantly influenced by thermal treatment, where Adelaide sharks raised at current temperatures had a lower capillarity than Jervis Bay sharks raised at ambient or projected EOC temperatures. This may indicate higher metabolic costs at elevated temperatures. However, our results suggest that regardless of the population, juvenile Port Jackson sharks may have limited acclimatory potential to alter muscle metabolic features under a temperature increase, which may make this species vulnerable to climate change.

Aquatic Walking and Swimming Kinematics of Neonate and Juvenile Epaulette Sharks.

The epaulette shark, Hemiscyllium ocellatum, is a small, reef-dwelling, benthic shark that-using its paired fins-can walk, both in and out of water. Within the reef flats, this species experiences short periods of elevated CO2 and hypoxia as well as fluctuating temperatures as reef flats become isolated with the outgoing tide. Past studies have shown that this species is robust (i.e., respiratory and metabolic performance, behavior) to climate change-relevant elevated CO2 levels as well as hypoxia and anoxia tolerant. However, epaulette shark embryos reared under ocean warming conditions hatch earlier and smaller, with altered patterns and coloration, and with higher metabolic costs than their current-day counterparts. Findings to date suggest that this species has adaptations to tolerate some, but perhaps not all, of the challenging conditions predicted for the 21st century. As such, the epaulette shark is emerging as a model system to understand vertebrate physiology in changing oceans. Yet, few studies have investigated the kinematics of walking and swimming, which may be vital to their biological fitness, considering their habitat and propensity for challenging environmental conditions. Given that neonates retain embryonic nutrition via an internalized yolk sac, resulting in a bulbous abdomen, while juveniles actively forage for worms, crustaceans, and small fishes, we hypothesized that difference in body shape over early ontogeny would affect locomotor performance. To test this, we examined neonate and juvenile locomotor kinematics during the three aquatic gaits they utilize-slow-to-medium walking, fast walking, and swimming-using 13 anatomical landmarks along the fins, girdles, and body midline. We found that differences in body shape did not alter kinematics between neonates and juveniles. Overall velocity, fin rotation, axial bending, and tail beat frequency and amplitude were consistent between early life stages. Data suggest that the locomotor kinematics are maintained between neonate and juvenile epaulette sharks, even as their feeding strategy changes. Studying epaulette shark locomotion allows us to understand this-and perhaps related-species' ability to move within and away from challenging conditions in their habitats. Such locomotor traits may not only be key to survival, in general, as a small, benthic mesopredator (i.e., movements required to maneuver into small reef crevices to avoid aerial and aquatic predators), but also be related to their sustained physiological performance under challenging environmental conditions, including those associated with climate change-a topic worthy of future investigation.

Species on the move around the Australian coastline: A continental-scale review of climate-driven species redistribution in marine systems.

Climate-driven changes in the distribution of species are a pervasive and accelerating impact of climate change, and despite increasing research effort in this rapidly emerging field, much remains unknown or poorly understood. We lack a holistic understanding of patterns and processes at local, regional and global scales, with detailed explorations of range shifts in the southern hemisphere particularly under-represented. Australian waters encompass the world's third largest marine jurisdiction, extending from tropical to sub-Antarctic climate zones, and have waters warming at rates twice the global average in the north and two to four times in the south. Here, we report the results of a multi-taxon continent-wide review describing observed and predicted species redistribution around the Australian coastline, and highlight critical gaps in knowledge impeding our understanding of, and response to, these considerable changes. Since range shifts were first reported in the region in 2003, 198 species from nine Phyla have been documented shifting their distribution, 87.3% of which are shifting poleward. However, there is little standardization of methods or metrics reported in observed or predicted shifts, and both are hindered by a lack of baseline data. Our results demonstrate the importance of historical data sets and underwater visual surveys, and also highlight that approximately one-fifth of studies incorporated citizen science. These findings emphasize the important role the public has had, and can continue to play, in understanding the impact of climate change. Most documented shifts are of coastal fish species in sub-tropical and temperate systems, while tropical systems in general were poorly explored. Moreover, most distributional changes are only described at the poleward boundary, with few studies considering changes at the warmer, equatorward range limit. Through identifying knowledge gaps and research limitations, this review highlights future opportunities for strategic research effort to improve the representation of Australian marine species and systems in climate-impact research.

Regulate or tolerate: Thermal strategy of a coral reef flat resident, the epaulette shark, Hemiscyllium ocellatum.

Highly variable thermal environments, such as coral reef flats, are challenging for marine ectotherms and are thought to invoke the use of behavioural strategies to avoid extreme temperatures and seek out thermal environments close to their preferred temperatures. Common to coral reef flats, the epaulette shark (Hemiscyllium ocellatum) possesses physiological adaptations to hypoxic and hypercapnic conditions, such as those experienced on reef flats, but little is known regarding the thermal strategies used by these sharks. We investigated whether H. ocellatum uses behavioural thermoregulation (i.e., movement to occupy thermally favourable microhabitats) or tolerates the broad range of temperatures experienced on the reef flat. Using an automated shuttlebox system, we determined the preferred temperature of H. ocellatum under controlled laboratory conditions and then compared this preferred temperature to 6 months of in situ environmental and body temperatures of individual H. ocellatum across the Heron Island reef flat. The preferred temperature of H. ocellatum under controlled conditions was 20.7 ± 1.5°C, but the body temperatures of individual H. ocellatum on the Heron Island reef flat mirrored environmental temperatures regardless of season or month. Despite substantial temporal variation in temperature on the Heron Island reef flat (15-34°C during 2017), there was a lack of spatial variation in temperature across the reef flat between sites or microhabitats. This limited spatial variation in temperature creates a low-quality thermal habitat limiting the ability of H. ocellatum to behaviourally thermoregulate. Behavioural thermoregulation is assumed in many shark species, but it appears that H. ocellatum may utilize other physiological strategies to cope with extreme temperature fluctuations on coral reef flats. While H. ocellatum appears to be able to tolerate acute exposure to temperatures well outside of their preferred temperature, it is unclear how this, and other, species will cope as temperatures continue to rise and approach their critical thermal limits. Understanding how species will respond to continued warming and the strategies they may use will be key to predicting future populations and assemblages.

Population variation in the thermal response to climate change reveals differing sensitivity in a benthic shark.

Many species with broad distributions are exposed to different thermal regimes which often select for varied phenotypes. This intraspecific variation is often overlooked but may be critical in dictating the vulnerability of different populations to environmental change. We reared Port Jackson shark (Heterodontus portusjacksoni) eggs from two thermally discrete populations (i.e. Jervis Bay and Adelaide) under each location's present-day mean temperatures, predicted end-of-century temperatures and under reciprocal-cross conditions to establish intraspecific thermal sensitivity. Rearing temperatures strongly influenced MO2 Max and critical thermal limits, regardless of population, indicative of acclimation processes. However, there were significant population-level effects, such that Jervis Bay sharks, regardless of rearing temperature, did not exhibit differences in MO2 Rest , but under elevated temperatures exhibited reduced maximum swimming activity with step-wise increases in temperature. In contrast, Adelaide sharks reared under elevated temperatures doubled their MO2 Rest , relative to their present-day temperature counterparts; however, maximum swimming activity was not influenced. With respect to reciprocal-cross comparisons, few differences were detected between Jervis Bay and Adelaide sharks reared under ambient Jervis Bay temperatures. Similarly, juveniles (from both populations) reared under Adelaide conditions had similar thermal limits and swimming activity (maximum volitional velocity and distance) to each other, indicative of conserved acclimation capacity. However, under Adelaide temperatures, the MO2 Rest of Jervis Bay sharks was greater than that of Adelaide sharks. This indicates that the energetics of cooler water population (Adelaide) is likely more thermally sensitive than that of the warmer population (Jervis Bay). While unique to elasmobranchs, these data provide further support that by treating species as static, homogeneous populations, we ignore the impacts of thermal history and intraspecific variation on thermal sensitivity. With climate change, intraspecific variation will manifest as populations move, demographics change or extirpations occur, starting with the most sensitive populations.

Bidirectional cyclical flows increase energetic costs of station holding for a labriform swimming fish, Cymatogaster aggregata.

Wave-induced surge conditions are found in shallow marine ecosystems worldwide; yet, few studies have quantified how cyclical surges may affect free swimming animals. Here, we used a recently adapted respirometry technique to compare the energetic costs of a temperate fish species (Cymatogaster aggregata) swimming against a steady flow versus cyclical unidirectional and bidirectional surges in which unsteady swimming (such as accelerating, decelerating and turning) occurs. Using oxygen uptake (MO2) as an estimate of energetic costs, our results reveal that fish swimming in an unsteady (i.e. cyclical) unidirectional flow showed no clear increase in costs when compared to a steady flow of the same average speed, suggesting that costs and savings from cyclical acceleration and coasting are near equal. Conversely, swimming in a bidirectional cyclical flow incurred significantly higher energetic costs relative to a steady, constant flow, likely due to the added cost of turning around to face the changing flow direction. On average, we observed a 50% increase in MO2 of fish station holding within the bidirectional flow (227.8 mg O2 kg-1 h-1) compared to a steady, constant flow (136.1 mg O2 kg-1 h-1) of the same mean velocity. Given wave-driven surge zones are prime fish habitats in the wild, we suggest the additional costs fish incur by station holding in a bidirectional cyclical flow must be offset by favourable conditions for foraging and reproduction. With current and future increases in abiotic stressors associated with climate change, we highlight the importance of incorporating additional costs associated with swimming in cyclical water flow in the construction of energy budgets for species living in dynamic, coastal habitats.

The emergence emergency: A mudskipper's response to temperatures.

Temperature has a profound effect on all life and a particularly influential effect on ectotherms, such as fishes. Amphibious fishes have a variety of strategies, both physiological and/or behavioural, to cope with a broad range of thermal conditions. This study examined the relationship between prolonged (5 weeks) exposure to a range of temperatures (22, 25, 28, or 32 °C) on oxygen uptake rate and movement behaviours (i.e., thermoregulation and emergence) in a common amphibious fish, the barred mudskipper (Periophthalmus argentilneatuis). At the highest temperature examined (32 °C, approximately 5 °C above their summer average temperatures), barred mudskippers exhibited 33.7-97.7% greater oxygen uptake rates at rest (MO2Rest), emerged at a higher temperature (CTe; i.e., a modified critical thermal maxima (CTMax) methodology) of 41.3 ±â€¯0.3 °C relative to those maintained at 28, 25, or 22 °C. The 32 °C-maintained fish also ceased movement activity at the highest holding temperature suggesting that prolonged submergence at elevated temperatures is physiologically and energetically stressful to the individual. Using exhaustive exercise protocols with and without air exposure to simulate a predatory chase, the time to recovery was examined for all individuals. When submerged, mudskippers required 2.5x longer recovery time to return to resting oxygen uptake from exhaustive exercise than those fully emerged in air. Oxygen uptake data revealed that air exposure did not accrue oxygen debt, thereby allowing faster return to resting oxygen consumption rates. If the option to emerge was not available, mudskippers preferentially sought more benign water temperatures (26.7 ±â€¯2.1 °C), resembling those experienced by these fish during the Austral autumn, regardless of prolonged exposure higher or lower temperatures. These results add to our understanding of the strategies that amphibious fishes may use to mitigate extra costs associated with living in warm waters, and could be the key to understanding how such species will cope with increasing temperatures in the future.