Hidden impacts of ocean warming and acidification on biological responses of marine animals revealed through meta-analysis.

Conflicting results remain on the impacts of climate change on marine organisms, hindering our capacity to predict the future state of marine ecosystems. To account for species-specific responses and for the ambiguous relation of most metrics to fitness, we develop a meta-analytical approach based on the deviation of responses from reference values (absolute change) to complement meta-analyses of directional (relative) changes in responses. Using this approach, we evaluate responses of fish and invertebrates to warming and acidification. We find that climate drivers induce directional changes in calcification, survival, and metabolism, and significant deviations in twice as many biological responses, including physiology, reproduction, behavior, and development. Widespread deviations of responses are detected even under moderate intensity levels of warming and acidification, while directional changes are mostly limited to more severe intensity levels. Because such deviations may result in ecological shifts impacting ecosystem structures and processes, our results suggest that climate change will likely have stronger impacts than those previously predicted based on directional changes alone.

The rostral micro-tooth morphology of blue marlin, Makaira nigricans.

Billfish rostra potentially have several functions; however, their role in feeding is unequivocal in some species. Recent work linked morphological variation in rostral micro-teeth to differences in feeding behavior in two billfish species, the striped marlin (Kajikia audax) and the sailfish (Istiophorus platypterus). Here, we present the rostral micro-tooth morphology for a third billfish species, the blue marlin (Makaira nigricans), for which the use of the rostrum in feeding behavior is still undocumented from systematic observations in the wild. We measured the micro-teeth on rostrum tips of blue marlin, striped marlin, and sailfish using a micro-computed tomography approach and compared the tooth morphology among the three species. This was done after an analysis of video-recorded hunting behavior of striped marlin and sailfish revealed that both species strike prey predominantly with the first third of the rostrum, which provided the justification to focus our analysis on the rostrum tips. In blue marlin, intact micro-teeth were longer compared to striped marlin but not to sailfish. Blue marlin had a higher fraction of broken teeth than both striped marlin and sailfish, and broken teeth were distributed more evenly on the rostrum. Micro-tooth regrowth was equally low in both marlin species but higher in sailfish. Based on the differences and similarities in the micro-tooth morphology between the billfish species, we discuss potential feeding-related rostrum use in blue marlin. We put forward the hypothesis that blue marlin might use their rostra in high-speed dashes as observed in striped marlin, rather than in the high-precision rostral strikes described for sailfish, possibly focusing on larger prey organisms.

Kinematics and behaviour in fish escape responses: guidelines for conducting, analysing and reporting experiments.

Work carried out since the late 1970s has provided key insights into the comparative biomechanics, kinematics, behaviour and neurobiology of fish escape responses. An escape response is an ecologically important behaviour used by fishes to evade predation and aggression via rapid swimming movements. With environmental change expected to affect the physiology and biomechanics of aquatic ectotherms, there is a growing interest in understanding how environmental stressors affect the swimming performance and behaviour of fishes during escape responses, particularly in the context of predator-prey interactions. As the study of fish swimming continues to expand, there have been repeated calls to standardise experiments and reporting practices to facilitate integrative and comparative studies. Here, we provide a set of practical guidelines for conducting, analysing and reporting experiments on escape responses in fish, including a reporting checklist to assist authors undertaking these experiments. These resources will facilitate executing and reporting escape response experiments in a rigorous and transparent fashion, helping to advance the study of fish swimming in an era of rapid environmental change.

Mechanisms of group-hunting in vertebrates.

Group-hunting is ubiquitous across animal taxa and has received considerable attention in the context of its functions. By contrast much less is known about the mechanisms by which grouping predators hunt their prey. This is primarily due to a lack of experimental manipulation alongside logistical difficulties quantifying the behaviour of multiple predators at high spatiotemporal resolution as they search, select, and capture wild prey. However, the use of new remote-sensing technologies and a broadening of the focal taxa beyond apex predators provides researchers with a great opportunity to discern accurately how multiple predators hunt together and not just whether doing so provides hunters with a per capita benefit. We incorporate many ideas from collective behaviour and locomotion throughout this review to make testable predictions for future researchers and pay particular attention to the role that computer simulation can play in a feedback loop with empirical data collection. Our review of the literature showed that the breadth of predator:prey size ratios among the taxa that can be considered to hunt as a group is very large (<100 to >102 ). We therefore synthesised the literature with respect to these predator:prey ratios and found that they promoted different hunting mechanisms. Additionally, these different hunting mechanisms are also related to particular stages of the hunt (search, selection, capture) and thus we structure our review in accordance with these two factors (stage of the hunt and predator:prey size ratio). We identify several novel group-hunting mechanisms which are largely untested, particularly under field conditions, and we also highlight a range of potential study organisms that are amenable to experimental testing of these mechanisms in connection with tracking technology. We believe that a combination of new hypotheses, study systems and methodological approaches should help push the field of group-hunting in new directions.

Multiple preferred escape trajectories are explained by a geometric model incorporating prey's turn and predator attack endpoint.

The escape trajectory (ET) of prey - measured as the angle relative to the predator's approach path - plays a major role in avoiding predation. Previous geometric models predict a single ET; however, many species show highly variable ETs with multiple preferred directions. Although such a high ET variability may confer unpredictability to avoid predation, the reasons why animals prefer specific multiple ETs remain unclear. Here, we constructed a novel geometric model that incorporates the time required for prey to turn and the predator's position at the end of its attack. The optimal ET was determined by maximizing the time difference of arrival at the edge of the safety zone between the prey and predator. By fitting the model to the experimental data of fish Pagrus major, we show that the model can clearly explain the observed multiple preferred ETs. By changing the parameters of the same model within a realistic range, we were able to produce various patterns of ETs empirically observed in other species (e.g., insects and frogs): a single preferred ET and multiple preferred ETs at small (20-50°) and large (150-180°) angles from the predator. Our results open new avenues of investigation for understanding how animals choose their ETs from behavioral and neurosensory perspectives.

When a prey spots a predator about to pounce, it turns swiftly and accelerates away to avoid being captured. The initial direction the prey chooses to take ­ known as its escape trajectory ­ can greatly impact their chance of survival. Previous models were able to predict the optimal direction an animal should take to maximize its chances of evading the predator. However, experimental data suggest that prey actually tend to escape via multiple specific directions, although why animals use this approach has not been clarified. To investigate this puzzle, Kawabata et al. built a new mathematical model that better represents how prey and predators interact with one another in the real world. Unlike past models, Kawabata et al. incorporated the time required for prey to change direction and only allowed the predators to move toward the prey for a limited distance. By including these two factors, they were able to reproduce the escape trajectories of real animals, including a species of fish, as well as species from other taxa such as frogs and insects. The new model suggests that prey escape along one of two directions: either by moving directly away from the predator in order to outrun its attack, or by dodging sideways to avoid being captured. Which strategy the prey chooses has some elements of unpredictability, which makes it more difficult for predators to adjust their capturing method. These findings shed light on why escaping in multiple specific directions makes prey harder to catch. The model could also be extended to test the escape trajectories of a wider variety of predator and prey species, which may avoid capture via different routes. This could help researchers better understand how predators and prey interact with one another. The findings could also reveal how sensory information (such as sound and sight) associated with the threat of an approaching predator is processed and stimulates the muscle activity required to escape in multiple different directions.

Combined effects of climate change stressors and predators with contrasting feeding-digestion strategies on a mussel species.

We investigated the combined effects of Ocean Warming (OW), Acidification (OA) and predator cues (Non-Consumptive Effects; NCEs) of two predators with contrasting feeding-digestion strategies on the mussel Perumytilus purpuratus. We considered starfish-NCEs (partially external digestion) and snail-NCEs (internal digestion). Mussels were exposed for 13 weeks to cross-factored OA (~500 and ~1400 µatm, pCO2) and OW (~15 and ~20 °C) conditions, in the presence/absence of NCEs from one or both predators. Mussels exposed to both NCEs exhibited smaller length and buoyant weight growth than those under control or snail-NCEs conditions. Mussels exposed to starfish-NCEs exhibited smaller wet mass than control mussels. OW and starfish-NCEs in isolation or combined with snail-NCEs increased mussel oxygen consumption. Byssal biogenesis was affected by the three-factors interaction. Clearance rates were affected by the OW × OA interaction. We suggest that mainly starfish-NCEs, in isolation or interacting with OA or/and OW, can threat mussel traits and the associated community.

How much biodiversity is concealed in the word 'biodiversity'?

Amidst a global biodiversity crisis1, the word 'biodiversity' has become indispensable for conservation and management2. Yet, biodiversity is often used as a buzzword in scientific literature. Resonant titles of papers claiming to have studied 'global biodiversity' may be used to promote research focused on a few taxonomic groups, habitats, or facets of biodiversity - taxonomic, (phylo)genetic, or functional. This usage may lead to extrapolating results outside the target systems of these studies with direct consequences for our understanding of life on Earth and its practical conservation. Here, we used a random sample of papers with the word 'biodiversity' in their title to take a long view of the use of this term. Despite improvements in analytical tools, monitoring technologies, and data availability3,4, we found that the taxonomic scope of research articles has not increased in recent years. We also show that studies with a wider taxonomic scope attract more citations and online attention. Our results have broad ramifications for understanding how extrapolating from studies with narrow taxonomic scope affects our view of global biodiversity and conservation.

Ocean acidification and seasonal temperature extremes combine to impair the thermal physiology of a sub-Antarctic fish.

To predict the potential impacts of climate change on marine organisms, it is critical to understand how multiple stressors constrain the physiology and distribution of species. We evaluated the effects of seasonal changes in seawater temperature and near-future ocean acidification (OA) on organismal and sub-organismal traits associated with the thermal performance of Eleginops maclovinus, a sub-Antarctic notothenioid species with economic importance to sport and artisanal fisheries in southern South America. Juveniles were exposed to mean winter and summer sea surface temperatures (4 and 10 °C) at present-day and near-future pCO2 levels (~500 and 1800 µatm). After a month, the Critical Thermal maximum and minimum (CTmax, CTmin) of fish were measured using the Critical Thermal Methodology and the aerobic scope of fish was measured based on the difference between their maximal and standard rates determined from intermittent flow respirometry. Lipid peroxidation and the antioxidant capacity were also quantified to estimate the oxidative damage potentially caused to gill and liver tissue. Although CTmax and CTmin were higher in individuals acclimated to summer versus winter temperatures, the increase in CTmax was minimal in juveniles exposed to the near-future compared to present-day pCO2 levels (there was a significant interaction between temperature and pCO2 on CTmax). The reduction in the thermal tolerance range under summer temperatures and near-future OA conditions was associated with a reduction in the aerobic scope observed at the elevated pCO2 level. Moreover, an oxidative stress condition was detected in the gill and liver tissues. Thus, chronic exposure to OA and the current summer temperatures pose limits to the thermal performance of juvenile E. maclovinus at the organismal and sub-organismal levels, making this species vulnerable to projected climate-driven warming.

Aerobic metabolic scope mapping of an invasive fish species with global warming.

Climate change will exacerbate the negative effects associated with the introduction of non-indigenous species in marine ecosystems. Predicting the spread of invasive species in relation to environmental warming is therefore a fundamental task in ecology and conservation. The Baltic Sea is currently threatened by several local stressors and the highest increase in sea surface temperature of the world's large marine ecosystems. These new thermal conditions can further favour the spreading of the invasive round goby (Neogobius melanostomus), a fish of Ponto-Caspian origin, currently well established in the southern and central parts of the Baltic Sea. This study aims to assess the thermal habitat suitability of the round goby in the Baltic Sea considering the past and future conditions. The study combines sightings records with known physiological models of aerobic performance and sea surface temperatures. Physiological models read these temperatures, at sighting times and locations, to determine their effects on the aerobic metabolic scope (AMS) of the fish, a measure of its energetic potential in relation to environmental conditions. The geographical mapping of the AMS was used to describe the changes in habitat suitability during the past 3 decades and for climatic predictions (until 2100) showing that the favourable thermal habitat in the Baltic Sea has increased during the past 32 years and will continue to do so in all the applied climate model predictions. Particularly, the predicted new thermal conditions do not cause any reduction in the AMS of round goby populations, while the wintertime cold ranges are likely expected to preserve substantial areas from invasion. The results of this research can guide future monitoring programs increasing the chance to detect this invader in novel areas.

Escape response kinematics in two species of tropical shark: short escape latencies and high turning performance.

Accelerative manoeuvres, such as fast-starts, are crucial for fish to avoid predation. Escape responses are fast-starts that include fundamental survival traits for prey that experience high predation pressure. However, no previous study has assessed escape performance in neonate tropical sharks. We quantitatively evaluated vulnerability traits of neonate tropical sharks by testing predictions on their fast-start escape performance. We predicted (1) high manoeuvrability, given their high flexibility, but (2) low propulsive locomotion owing to the drag costs associated with pectoral fin extension during escape responses. Further, based on previous work on dogfish, Squalus suckleyi, we predicted (3) long reaction times (as latencies longer than teleosts, >20 ms). We used two-dimensional, high-speed videography analysis of mechano-acoustically stimulated neonate blacktip reef shark, Carcharhinus melanopterus (n=12), and sicklefin lemon shark, Negaprion acutidens (n=8). Both species performed a characteristic C-start double-bend response (i.e. two body bends), but single-bend responses were only observed in N. acutidens. As predicted, neonate sharks showed high manoeuvrability with high turning rates and tight turning radii (3-11% of body length) but low propulsive performance (i.e. speed, acceleration and velocity) when compared with similar-sized teleosts and S. suckleyi. Contrary to expectations, escape latencies were <20 ms in both species, suggesting that the neurophysiological system of sharks when reacting to a predatory attack may not be limited to long response times. These results provide a quantitative assessment of survival traits in neonate tropical sharks that will be crucial for future studies that consider the vulnerability of these sharks to predation.

Escaping from multiple visual threats: modulation of escape responses in Pacific staghorn sculpin (Leptocottus armatus).

Fish perform rapid escape responses to avoid sudden predatory attacks. During escape responses, fish bend their bodies into a C-shape and quickly turn away from the predator and accelerate. The escape trajectory is determined by the initial turn (stage 1) and a contralateral bend (stage 2). Previous studies have used a single threat or model predator as a stimulus. In nature, however, multiple predators may attack from different directions simultaneously or in close succession. It is unknown whether fish are able to change the course of their escape response when startled by multiple stimuli at various time intervals. Pacific staghorn sculpin (Leptocottus armatus) were startled with a left and right visual stimulus in close succession. By varying the timing of the second stimulus, we were able to determine when and how a second stimulus could affect the escape response direction. Four treatments were used: a single visual stimulus (control); or two stimuli coming from opposite sides separated by a 0 ms (simultaneous treatment), 33 ms or 83 ms time interval. The 33 ms and 83 ms time intervals were chosen to occur either side of a predicted 60 ms visual escape latency (i.e. during stage 1). The 0 ms and 33 ms treatments influenced both the escape trajectory and the stage 1 turning angle, compared with a single stimulation, whereas the 83 ms treatment had no effect on the escape trajectory. We conclude that Pacific staghorn sculpin can modulate their escape trajectory only between stimulation and the onset of the response, but the escape trajectory cannot be modulated after the body motion has started.

Multiple-stressor effects of ocean acidification, warming and predation risk cues on the early ontogeny of a rocky-shore keystone gastropod.

To understand how climate change stressors might affect marine organisms and support adequate projections it is important to know how multiple stressors may be modulated by the presence of other species. We evaluated the direct effects of ocean warming (OW) and ocean acidification (OA) together with non-consumptive effects (NCEs) of the predatory crab Acanthocyclus hassleri on early ontogeny fitness-related traits of the commercially important rocky-shore keystone gastropod Concholepas concholepas. We measured the response of nine traits to these stressors at either the organismal level (survival, growth, feeding rates, tenacity, metabolic rate, calcification rate) or sub-organismal level (nutritional status, ATP-supplying capacity, stress condition). C. concholepas survival was not affected by any of the stressors. Feeding rates were not affected by OW or OA; however, they were reduced in the presence of crab NCEs compared with control conditions. Horizontal tenacity was affected by the OA × NCEs interaction; in the presence of NCEs, OA reduced tenacity. The routine metabolic rate, measured by oxygen consumption, increased significantly with OW. Nutritional status assessment determined that carbohydrate content was not affected by any of the stressors. However, protein content was affected by the OA × NCEs interaction; in the absence of NCEs, OA reduced protein levels. ATP-supplying capacity, measured by citrate synthase (CS) activity, and cellular stress condition (HSP70 expression) were reduced by OA, with reduction in CS activity found particularly at the high temperature. Our results indicate C. concholepas traits are affected by OA and OW and the effects are modulated by predator risk (NCEs). We conclude that some C. concholepas traits are resilient to climate stressors (survival, growth, horizontal tenacity and nutritional status) but others are affected by OW (metabolic rate), OA (ATP-supplying capacity, stress condition), and NCEs (feeding rate). The results suggest that these negative effects can adversely affect the associated community.

Lacunae rostralis: A new structure on the rostrum of sailfish Istiophorus platypterus.

Recent comparative studies of billfishes (Istiophoridae and Xiphiidae) have provided evidence of differences in the form and function of the rostra (bill) among species. Here, we report the discovery of a new structure, lacuna rostralis, on the rostra of sailfish Istiophorus platypterus, which is absent on the rostra of swordfish Xiphias gladius, striped marlin Kajikia audax and blue marlin Makaira nigricans. The lacunae rostralis are small cavities that contain teeth. They were found on the ventral rostrum surface of all I. platypterus specimens examined and dorsally in half of them. Ventrally, the lacunae rostralis were most prominent in the mid-section of the rostrum. Dorsally, they occurred closer to the tip. The density of lacunae rostralis increased towards the rostrum tip but, because they are smaller in size, the percentage of rostrum coverage decreased. The teeth located within the lacunae rostralis were found to be different in size, location and orientation from the previously identified micro-teeth of billfish. We propose two potential functions of the lacunae rostralis that both relate to the use of the bill in feeding: mechanoreception of prey before tapping it with the bill and more efficient prey handling via the creation of suction, or physical grip.

Social familiarity improves fast-start escape performance in schooling fish.

Using social groups (i.e. schools) of the tropical damselfish Chromis viridis, we test how familiarity through repeated social interactions influences fast-start responses, the primary defensive behaviour in a range of taxa, including fish, sharks, and larval amphibians. We focus on reactivity through response latency and kinematic performance (i.e. agility and propulsion) following a simulated predator attack, while distinguishing between first and subsequent responders (direct response to stimulation versus response triggered by integrated direct and social stimulation, respectively). In familiar schools, first and subsequent responders exhibit shorter latency than unfamiliar individuals, demonstrating that familiarity increases reactivity to direct and, potentially, social stimulation. Further, familiarity modulates kinematic performance in subsequent responders, demonstrated by increased agility and propulsion. These findings demonstrate that the benefits of social recognition and memory may enhance individual fitness through greater survival of predator attacks.

Age and sex influence social interactions, but not associations, within a killer whale pod.

Social structure is a fundamental aspect of animal populations. In order to understand the function and evolution of animal societies, it is important to quantify how individual attributes, such as age and sex, shape social relationships. Detecting these influences in wild populations under natural conditions can be challenging, especially when social interactions are difficult to observe and broad-scale measures of association are used as a proxy. In this study, we use unoccupied aerial systems to observe association, synchronous surfacing, and physical contact within a pod of southern resident killer whales (Orcinus orca). We show that interactions do not occur randomly between associated individuals, and that interaction types are not interchangeable. While age and sex did not detectably influence association network structure, both interaction networks showed significant social homophily by age and sex, and centrality within the contact network was higher among females and young individuals. These results suggest killer whales exhibit interesting parallels in social bond formation and social life histories with primates and other terrestrial social mammals, and demonstrate how important patterns can be missed when using associations as a proxy for interactions in animal social network studies.

The combined effects of climate change stressors and predatory cues on a mussel species.

In order to make adequate projections on the consequences of climate change stressors on marine organisms, it is important to know how impacts of these stressors are affected by the presence of other species. Here we assessed the direct effects of ocean warming (OW) and acidification (OA) along with non-consumptive effects (NCEs) of a predatory crab and/or a predatory snail on the habitat-forming mussel Perumytilus purpuratus. Mussels were exposed for 10-14 weeks to contrasting pCO2 (500 and 1400 µatm) and temperature (15 and 20 °C) levels, in the presence/absence of cues from one or two predator species. We compared mussel traits at sub-organismal (nutritional status, metabolic capacity-ATP production-, cell stress condition via HSP70 expression) and organismal (survival, oxygen consumption, growth, byssus biogenesis, clearance rates, aggregation) levels. OA increased the mussels' oxygen consumption; and OA combined with OW increased ATP demand and the use of carbohydrate reserves. Mussels at present-day pCO2 levels had the highest protein content. Under OW the predatory snail cues induced the highest cell stress condition on the mussels. Temperature, predator cues and the interaction between them affected mussel growth. Mussels grew larger at the control temperature (15 °C) when crab and snail cues were present. Mussel wet mass and calcification were affected by predator cues; with highest values recorded in crab cue presence (isolated or combined with snail cues). In the absence of predator cues in the trails, byssus biogenesis was affected by OA, OW and the OA × OW and OA × predator cues interactions. At present-day pCO2 levels, more byssus was recorded with snail than with crab cues. Clearance rates were affected by temperature, pCO2 and the interaction between them. The investigated stressors had no effects on mussel aggregation. We conclude that OA, OW and the NCEs may lead to neutral, positive or negative consequences for mussels.

Latency of mechanically stimulated escape responses in the Pacific spiny dogfish, Squalus suckleyi.

Fast escape responses to a predator threat are fundamental to the survival of mobile marine organisms. However, elasmobranchs are often underrepresented in such studies. Here, we measured the escape latency (time interval between the stimulus and first visible reaction) of mechanically induced escape responses in the Pacific spiny dogfish, Squalus suckleyi, and in two teleosts from the same region, the great sculpin, Myoxocephalus polyacanthocephalus, and the pile perch, Rhacochilus vacca We found that the dogfish had a longer minimum latency (66.7 ms) compared with that for the great sculpin (20.8 ms) and pile perch (16.7 ms). Furthermore, the dogfish had a longer latency than that of 48 different teleosts identified from 35 different studies. We suggest such long latencies in dogfish may be due to the absence of Mauthner cells, the giant neurons that control fast escape responses in fishes.

The combined effects of ocean acidification and warming on a habitat-forming shell-crushing predatory crab.

In mid rocky intertidal habitats the mussel Perumytilus purpurarus monopolizes the substratum to the detriment of many other species. However, the consumption of mussels by the shell-crushing crab Acanthocyclus hassleri creates within the mussel beds space and habitat for several other species. This crab uses its disproportionately large claw to crush its shelled prey and plays an important role in maintaining species biodiversity. This study evaluated the consequences of projected near-future ocean acidification (OA) and warming (OW) on traits of A. hassleri linked with their predatory performance. Individual A. hassleri were maintained for 10-16 weeks under contrasting pCO2 (~500 and 1400 µatm) and temperature (~15 and 20 °C) levels. We compared traits at the organismal (oxygen consumption rate, survival, calcification rate, feeding rates, crusher claw pinching strength, self-righting speed, sarcomere length of the crusher claw muscles) and cellular (nutritional status ATP provisioning capacity through citrate synthase activity, expression of HSP70) level. Survival, calcification rate and sarcomere length were not affected by OA and OW. However, OW increased significantly feeding and oxygen consumption. Pinching strength was reduced by OA; meanwhile self-righting was increased by OA and OW. At 20 °C, carbohydrate content was reduced significantly by OA. Regardless of temperature, a significant reduction in energy reserves in terms of protein content by OA was found. The ATP provisioning capacity was significantly affected by the interaction between temperature and pCO2 and was highest at 15 °C and present day pCO2 levels. The HSP70 levels of crabs exposed to OW were higher than in the control crabs. We conclude that OA and OW might affect the amount and size of prey consumed by this crab. Therefore, by reducing the crab feeding performance these stressors might pose limits on their role in generating microhabitat for other rocky intertidal species inhabiting within mussel beds.

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.