Acoustic scaling in the European spiny lobster (Palinurus elephas).

Sound is an important cue for arthropods. In insects, sound features and sound-producing apparatus are tightly correlated to enhance signal emission in larger individuals. In contrast, acoustic scaling in marine arthropods is poorly described even if they possess similar sound-producing apparatus. Here, the acoustic scaling of the European spiny lobster is analyzed by recording sounds in situ at 1 m from a wide range of body sizes. The dimensions of associated sound-producing apparatus increased with body size, indicating sound features would also be influenced by spiny lobster size. Indeed, temporal sound features changed with body size, suggesting differences in calling songs could be used for spiny lobster acoustic communication. Source levels (peak-peak) ranged from 131 to 164 dB re 1µPa for smaller and larger lobsters, respectively, which could be explained by more efficient resonating structures in larger animals. In addition, dominant frequencies were highly constrained by ambient noise levels, masking the low-frequency content of low intensity sounds from smaller spiny lobsters. Although the ecological function of spiny lobster sounds is not clear yet, these results suggest larger body sizes benefit because louder calls increase the broadcast area and potential interactions with conspecifics, as shown in the insect bioacoustic literature.

Insights into the behavioural responses of juvenile thornback ray Raja clavata to alternating and direct current magnetic fields.

As part of energy transition, marine renewable energy devices (MRED) are currently expanding in developed countries inducing the deployment of dense networks of submarine power cables. Concern has thus raised about the cable magnetic emissions (direct or alternating current) because of potential interference with the sensorial environment of magneto-sensitive species, such as sharks and rays. This study sought to assess the short-term behavioural responses of juvenile thornback rays (Raja clavata) (n = 15) to direct and alternating (50 Hz) uniform 450-µT artificial magnetic fields using 1 h focal-sampling design based on a detailed ethogram. Careful control of magnetic fields' temporal and spatial scales was obtained in laboratory conditions through a custom-made Helmholtz coil device. Overall, qualitative or quantitative behavioural responses of juvenile rays did not significantly vary between control vs. exposed individuals over the morning period. Nonetheless, rays under direct current magnetic field increased their activity over the midday period. Synchronisation patterns were also observed for individuals receiving alternating current exposure (chronologic and qualitative similarities) coupled with a high inter-individual variance. Further studies should consider larger batches of juveniles to address the effect of long-term exposure and explore the sensitivity range of rays with dose-response designs.

Sound detection by the American lobster (Homarus americanus).

Although many crustaceans produce sounds, their hearing abilities and mechanisms are poorly understood, leaving uncertainties regarding whether or how these animals use sound for acoustic communication. Marine invertebrates lack gas-filled organs required for sound pressure detection, but some of them are known to be sensitive to particle motion. Here, we examined whether the American lobster (Homarus americanus) could detect sound and subsequently sought to discern the auditory mechanisms. Acoustic stimuli responses were measured using auditory evoked potential (AEP) methods. Neurophysiological responses were obtained from the brain using tone pips between 80 and 250 Hz, with best sensitivity at 80-120 Hz. There were no significant differences between the auditory thresholds of males and females. Repeated controls (recordings from deceased lobsters, moving electrodes away from the brain and reducing seawater temperature) indicated the evoked potentials' neuronal origin. In addition, AEP responses were similar before and after antennules (including statocysts) were ablated, demonstrating that the statocysts, a long-proposed auditory structure in crustaceans, are not the sensory organs responsible for lobster sound detection. However, AEPs could be eliminated (or highly reduced) after immobilizing hairfans, which cover much of lobster bodies. These results suggest that these external cuticular hairs are likely to be responsible for sound detection, and imply that hearing is mechanistically possible in a wider array of invertebrates than previously considered. Because the lobsters' hearing range encompasses the fundamental frequency of their buzzing sounds, it is likely that they use sound for intraspecific communication, broadening our understanding of the sensory ecology of this commercially vital species. The lobsters' low-frequency acoustic sensitivity also underscores clear concerns about the potential impacts of anthropogenic noise.

Global COVID-19 lockdown highlights humans as both threats and custodians of the environment.

The global lockdown to mitigate COVID-19 pandemic health risks has altered human interactions with nature. Here, we report immediate impacts of changes in human activities on wildlife and environmental threats during the early lockdown months of 2020, based on 877 qualitative reports and 332 quantitative assessments from 89 different studies. Hundreds of reports of unusual species observations from around the world suggest that animals quickly responded to the reductions in human presence. However, negative effects of lockdown on conservation also emerged, as confinement resulted in some park officials being unable to perform conservation, restoration and enforcement tasks, resulting in local increases in illegal activities such as hunting. Overall, there is a complex mixture of positive and negative effects of the pandemic lockdown on nature, all of which have the potential to lead to cascading responses which in turn impact wildlife and nature conservation. While the net effect of the lockdown will need to be assessed over years as data becomes available and persistent effects emerge, immediate responses were detected across the world. Thus, initial qualitative and quantitative data arising from this serendipitous global quasi-experimental perturbation highlights the dual role that humans play in threatening and protecting species and ecosystems. Pathways to favorably tilt this delicate balance include reducing impacts and increasing conservation effectiveness.

Acoustic behaviour of male European lobsters (Homarus gammarus) during agonistic encounters.

Previous studies have demonstrated that male European lobsters (Homarus gammarus) use chemical and visual signals as a means of intraspecific communication during agonistic encounters. In this study, we show that they also produce buzzing sounds during these encounters. This result was missed in earlier studies because low-frequency buzzing sounds are highly attenuated in tanks, and are thus difficult to detect with hydrophones. To address this issue, we designed a behavioural tank experiment using hydrophones, with accelerometers placed on the lobsters to directly detect their carapace vibrations (i.e. the sources of the buzzing sounds). While we found that both dominant and submissive individuals produced carapace vibrations during every agonistic encounter, very few of the associated buzzing sounds (15%) were recorded by the hydrophones. This difference is explained by their high attenuation in tanks. We then used the method of algorithmic complexity to analyse the carapace vibration sequences as call-and-response signals between dominant and submissive individuals. Even though some intriguing patterns appeared for closely size-matched pairs (<5 mm carapace length difference), the results of the analysis did not permit us to infer that the processes underlying these sequences could be differentiated from random ones. Thus, such results prevented any conclusions about acoustic communication. This concurs with both the high attenuation of the buzzing sounds during the experiments and the poor understanding of acoustic perception by lobsters. New approaches that circumvent tank acoustic issues are now required to validate the existence of acoustic communication in lobsters.

Shells of the bivalve Astarte moerchi give new evidence of a strong pelagic-benthic coupling shift occurring since the late 1970s in the North Water polynya.

Climate changes in the Arctic may weaken the currently tight pelagic-benthic coupling. In response to decreasing sea ice cover, arctic marine systems are expected to shift from a 'sea-ice algae-benthos' to a 'phytoplankton-zooplankton' dominance. We used mollusc shells as bioarchives and fatty acid trophic markers to estimate the effects of the reduction of sea ice cover on the food exported to the seafloor. Bathyal bivalve Astarte moerchi living at 600 m depth in northern Baffin Bay reveals a clear shift in growth variations and Ba/Ca ratios since the late 1970s, which we relate to a change in food availability. Tissue fatty acid compositions show that this species feeds mainly on microalgae exported from the euphotic zone to the seabed. We, therefore, suggest that changes in pelagic-benthic coupling are likely due either to local changes in sea ice dynamics, mediated through bottom-up regulation exerted by sea ice on phytoplankton production, or to a mismatch between phytoplankton bloom and zooplankton grazing due to phenological change. Both possibilities allow a more regular and increased transfer of food to the seabed. This article is part of the theme issue 'The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'.

Human activities and climate variability drive fast-paced change across the world's estuarine-coastal ecosystems.

Time series of environmental measurements are essential for detecting, measuring and understanding changes in the Earth system and its biological communities. Observational series have accumulated over the past 2-5 decades from measurements across the world's estuaries, bays, lagoons, inland seas and shelf waters influenced by runoff. We synthesize information contained in these time series to develop a global view of changes occurring in marine systems influenced by connectivity to land. Our review is organized around four themes: (i) human activities as drivers of change; (ii) variability of the climate system as a driver of change; (iii) successes, disappointments and challenges of managing change at the sea-land interface; and (iv) discoveries made from observations over time. Multidecadal time series reveal that many of the world's estuarine-coastal ecosystems are in a continuing state of change, and the pace of change is faster than we could have imagined a decade ago. Some have been transformed into novel ecosystems with habitats, biogeochemistry and biological communities outside the natural range of variability. Change takes many forms including linear and nonlinear trends, abrupt state changes and oscillations. The challenge of managing change is daunting in the coastal zone where diverse human pressures are concentrated and intersect with different responses to climate variability over land and over ocean basins. The pace of change in estuarine-coastal ecosystems will likely accelerate as the human population and economies continue to grow and as global climate change accelerates. Wise stewardship of the resources upon which we depend is critically dependent upon a continuing flow of information from observations to measure, understand and anticipate future changes along the world's coastlines.

Noise pollution causes parental stress on marine invertebrates, the Giant scallop example.

In marine invertebrates, abiotic stresses on adults can act directly on gametes quality, which impacts phenotype and development success of the offspring. Human activities introduce noise pollution in the marine environment but still few studies on invertebrates have considered the impacts on adult or larval stages separately, and to our knowledge, never investigated the cross-generational effects of anthropogenic noise. This article explores parental effects of pile driving noise associated with the building phase of offshore wind turbines on a coastal invertebrate, Pecten maximus (L.). Adults were exposed to increasing levels of sound during gametogenesis, then their offspring were also exposed. The results highlight that anthropogenic noise experienced by the parents reduces their reproductive investment and modify larval response in similar conditions. Also, larvae from exposed adults grew 6-fold faster and metamorphosed 5-fold faster, which could be an amplified adaptive strategy to reduce the pelagic phase in a stressful environment.

Offshore windfarm construction elevates metabolic rate and increases predation vulnerability of a key marine invertebrate.

A global increase in offshore windfarm development is critical to our renewable energy future. Yet, widespread construction plans have generated substantial concern for impacts to co-occurring organisms and the communities they form. Pile driving construction, prominent in offshore windfarm development, produces among the highest amplitude sounds in the ocean creating widespread concern for a diverse array of taxa. However, studies addressing ecologically key species are generally lacking and most research is disparate, failing to integrate across response types (e.g., behavior, physiology, and ecological interactions), particularly in situ. The lack of integrative field studies presents major challenges to understand or mitigate actual impacts of offshore wind development. Here, we examined critical behavioral, physiological, and antipredator impacts of actual pile driving construction on the giant sea scallop (Placopecten magellanicus). Benthic taxa including bivalves are of particular concern because they are sound-sensitive, cannot move appreciable distances away from the stressor, and support livelihoods as one of the world's most economically and socially important fisheries. Overall, pile driving sound impacted scallops across a series of behavioral and physiological assays. Sound-exposed scallops consistently reduced their valve opening (22%), resulting in lowered mantle water oxygen levels available to the gills. Repeated and rapid valve adductions led to a 56% increase in metabolic rates relative to pre-exposure baselines. Consequently, in response to predator stimuli, sound-exposed scallops displayed a suite of significantly weaker antipredator behaviors including fewer swimming events and shorter time-to-exhaustion. These results show aquatic construction activities can induce metabolic and ecologically relevant changes in a key benthic animal. As offshore windfarm construction accelerates globally, our field-based study highlights that spatial overlap with benthic taxa may cause substantial metabolic changes, alter important fisheries resources, and ultimately could lead to increased predation.

Effects of anthropogenic magnetic fields on the behavior of a major predator of the intertidal and subtidal zones, the velvet crab Necora puber.

With the progress of the offshore renewable energy sector and electrical interconnection projects, a substantial rise in the number of submarine power cables is expected soon. Such cables emit either alternating or direct current magnetic fields whose impact on marine invertebrates is currently unknown and hardly studied. In this context, this study aimed to assess potential short-term exposure (30 min) effects of both alternating and direct magnetic fields of increasing intensity (72-304 µT) on the behavior of the high-ecological value velvet crab (Necora puber). Three experiments were designed to evaluate whether the strongest magnetic field intensities induce crabs' attraction or repulsion responses, and whether foraging and sheltering behaviors may be modified. We extracted from video analyses several variables as the time budgets crabs spent immobile, moving, feeding, or sheltering as well as total and maximal distance reached in the magnetic field (MF) gradient. The crabs exposed to artificial MF did not exhibit significant behavioral changes compared with those exposed to the "natural" MF. Overall, our results suggest that, at such intensities, artificial magnetic fields do not significantly alter behaviors of N. puber. Nevertheless, future studies should be conducted to examine the effects of longer exposure periods and to detect potential habituation or resilience processes.

Influence of sea-ice dynamics on coastal Antarctic benthos: A case study on lantern clams (Laternula elliptica) in Adélie Land.

Polar regions are warming faster than the world average and are profoundly affected by changes in the spatio-temporal dynamics of sea ice, with largely unknown repercussions on the functioning of marine ecosystems. Here, we investigated the impacts of interannual sea-ice variability on coastal benthic communities in Antarctica, focusing on a close-to-pristine area (Adélie Land). We investigated shell growth of the circum-Antarctic bivalve Laternula elliptica, considered a key species in these soft bottom benthic communities. Chondrophores of live-collected clams were prepared using standard sclerochronological methods to study the interannual variability of shell growth from 1996 to 2015. Our results show that the master chronology varied with sea-ice dynamics. When sea ice breaks up too early, sympagic algae do not have time to accumulate sufficiently high biomass, thus strongly limiting the energy input to the benthos. This negatively affects the physiological performance of L. elliptica, thereby altering their population dynamics and hence the functioning of these soft-bottom ecosystems.

Three-dimensional anthropogenic underwater noise modeling in an Arctic fjord for acoustic risk assessment.

Sea-ice covering is drastically declining in the Arctic, opening new maritime routes and thus introducing underwater noise pollution in nearly pristine acoustic environments. Evaluating underwater noise pollution requires good acoustic propagation modeling to predict sound exposure levels. However, underwater noise modeling for acoustic risk assessments has often been carried out using simplistic propagation models, which approximate a 3D propagation in several planes (Nx2D), instead of using full 3D propagation models. However, Nx2D propagation models are impractical for winding geography and steep bathymetry as found in Arctic fjords. The purpose of this study is to estimate disturbance and masking effects on Arctic animal species from shipping noises, modeled through a traditional Nx2D BELLHOP model and a full 3D BELLHOP model. Classical Nx2D propagation modeling largely underestimates the anthropogenic noise footprint in Arctic fjords compared to using a full 3D propagation model.

The nocturnal life of the great scallops (Pecten maximus, L.): First description of their natural daily valve opening cycle.

Valvometry techniques used to monitor bivalve gaping activity have elucidated numerous relationships with environmental fluctuations, along with biological rhythms ranging from sub-daily to seasonal. Thus, a precise understanding of the natural activity of bivalves (i.e., not exposed to stressful environmental variations) is necessary as a baseline for detecting abnormal behaviors (deviations). This knowledge is also needed to reliably interpret observations of bivalve gaping behavior and associated biological processes (e.g., respiration, nutrition) acquired over time-limited periods. With this in mind, we investigated the natural daily gaping activity of the great scallop (Pecten maximus) by continuously monitoring 35 individuals in several individual tanks and in situ (Bay of Saint-Brieuc, Brittany, France) using fully autonomous Hall effect sensors. Our results revealed a circadian cycle (τ = 24.0h) in scallop gaping activity. Despite significant inter-individual variability in mean opening and cycle amplitude, almost all individuals (87.5%) exhibited nocturnal activity, with valves more open at night than during the day. A shift in light regime in the tanks triggered an instantaneous change in opening pattern, indicating that light levels strongly determine scallop activity. Based on the opening status of scallops, we also identified several gaping behaviors deviating from the regular daily pattern (lack of rhythmicity, high daytime opening), potentially reflecting physiological weakness. While further long-term studies are required to fully understand the natural activity of scallops, these findings pave the way for studies focused on the scallop response to external factors and introduce further research into the detection of abnormal behaviors. Coupling observations of diel valve gaping cycles with other daily variations in organismal and environmental parameters could help explain mechanisms driving the growth patterns of scallops observed in their shell striations. From a technical perspective, our field-based monitoring demonstrates the suitability of autonomous valvometry sensors for studying mobile subtidal bivalve activity in remote offshore environments.

Seasonal lipid dynamics of four Arctic bivalves: Implications for their physiological capacities to cope with future changes in coastal ecosystems.

The Arctic is exposed to unprecedented warming, at least three times higher than the global average, which induces significant melting of the cryosphere. Freshwater inputs from melting glaciers will subsequently affect coastal primary production and organic matter quality. However, due to a lack of basic knowledge on the physiology of Arctic organisms, it remains difficult to understand how these future trophic changes will threaten the long-term survival of benthic species in coastal habitats. This study aimed to gain new insights into the seasonal lipid dynamics of four dominant benthic bivalves (Astarte moerchi, Hiatella arctica, Musculus discors, and Mya truncata) collected before and after sea ice break-up in a high-Arctic fjord (Young Sound, NE Greenland). Total lipid content and fatty acid composition of digestive gland neutral lipids were analyzed to assess bivalve energy reserves while the fatty acid composition of gill polar lipids was determined as a biochemical indicator of interspecies variations in metabolic activity and temperature acclimation. Results showed a decrease in lipid reserves between May and August, suggesting that bivalves have only limited access to fresh organic matter until sea ice break-up. The lack of seasonal variation in the fatty acid composition of neutral lipids, especially essential ω3 fatty acids, indicates that no fatty acid transfer from the digestive glands to the gonads occurs between May and August, and therefore, no reproductive investment takes place during this period. Large interspecies differences in gill fatty acid composition were observed, which appear to be related to differences in species life span and metabolic strategies. Such differences in gill fatty acid composition of polar lipids, which generally influence metabolic rates and energy needs, may imply that not all benthic species will be equally sensitive to future changes in primary production and organic matter quality in Arctic coastal habitats.

Pile driving and drilling underwater sounds impact the metamorphosis dynamics of Pecten maximus (L., 1758) larvae.

One of the biggest challenges of the 21st century is to reduce carbon emissions and offshore wind turbines seem to be an efficient solution. However, during the installation phase, high levels of noise are emitted whose impacts remain not well known, particularly on benthic marine invertebrates displaying a bentho-planktonic life-cycle. For one century, larval settlement and subsequent recruitment has been considered as a key topic in ecology as it determines largely population renewal. Whereas several recent studies have shown that trophic pelagic but also natural soundscape cues could trigger bivalve settlement, the role of anthropogenic noise remains poorly documented. Therefore, we conducted experiments to assess potential interacting effects of diet and pile driving or drilling sounds on the great scallop (Pecten maximus) larval settlement. We demonstrate here that pile driving noise stimulates both growth and metamorphosis as well as it increases the total lipid content of competent larvae. Conversely, drilling noise reduces both survival and metamorphosis rates. For the first time, we provide evidence of noise impacts associated to MREs installation on P. maximus larvae and discuss about potential consequences on their recruitment.

Potential for acoustic masking due to shipping noise in the European lobster (Homarus gammarus).

Marine traffic is the most pervasive underwater anthropogenic noise pollution which can mask acoustic communication in marine mammals and fish, but its effect in marine invertebrates remains unknown. Here, we performed an at sea experiment to study the potential of shipping noise to mask and alter lobster acoustic communication. We used hydrophones to record buzzing sounds and accelerometers to detect lobster carapace vibrations (i.e. the buzzing sounds' sources). We demonstrated that male individuals produced carapace vibrations under various ambient noise conditions, including heavy shipping noise. However, while the associated waterborne buzzing sounds could be recorded under natural ambient noise levels, they were masked by shipping noise. Additionally, lobsters significantly increased their call rates in presence of shipping noise, suggesting a vocal compensation due to the reduction of intraspecific communication. This study reports for the first time the potential acoustic masking of lobster acoustic communication by chronic anthropogenic noise pollution, which could affect ecologically important behaviors.

Spiny lobster sounds can be detectable over kilometres underwater.

The detection ranges of broadband sounds produced by marine invertebrates are not known. To address this deficiency, a linear array of hydrophones was built in a shallow water area to experimentally investigate the propagation features of the sounds from various sizes of European spiny lobsters (Palinurus elephas), recorded between 0.5 and 100 m from the animals. The peak-to-peak source levels (SL, measured at one meter from the animals) varied significantly with body size, the largest spiny lobsters producing SL up to 167 dB re 1 µPa2. The sound propagation and its attenuation with the distance were quantified using the array. This permitted estimation of the detection ranges of spiny lobster sounds. Under the high ambient noise conditions recorded in this study, the sounds propagated between 5 and 410 m for the smallest and largest spiny lobsters, respectively. Considering lower ambient noise levels and different realistic propagation conditions, spiny lobster sounds can be detectable up to several kilometres away from the animals, with sounds from the largest individuals propagating over 3 km. Our results demonstrate that sounds produced by P. elephas can be utilized in passive acoustic programs to monitor and survey this vulnerable species at kilometre scale in coastal waters.

Does trace element composition of bivalve shells record utra-high frequency environmental variations?

Saint-Pierre and Miquelon (SPM) is a small archipelago where instrumental measures based on water column velocity and temperature profiles compiled comprehensive evidence for strong near-diurnal (25.8h) current and bottom temperature oscillations (up to 11.5 °C) which is possibly the largest ever observed - at any frequency - on a stratified mid-latitude continental shelf. The main objective of our study was to identify if Placopecten magellanicus can record on its shell these high frequency environmental variations. To this end, we have tried to identify proxies for water temperature and food availability through development of a new ultra-high resolution LA-ICPMS analyses method capable of resolving shell surface elemental composition with a 10 µm resolution. This method was applied on two shell fragments, both representing the third year of growth and 2015 annual growth period, respectively coming from two environmentally contrasted sites, more (30 m depth) or less (10 m depth) affected by high frequency thermal oscillations. Our results strongly suggest a relationship between phytoplankton biomass and barium incorporation into P. magellanicus shells at both sites. Even if P. magellanicus might present a physiological control of magnesium incorporation, the shape of the two Mg/Ca profiles seems to illustrate that temperature also exerts a control on magnesium incorporation in P. magellanicus shells from SPM. While U/Ca and Mg/Ca profiles show a strong positive correlation for 30 m site shell, suggesting that uranium incorporation in P. magellanicus shell is at least partially temperature dependent. The absence of such correlation for 10 m site shell suggests differences in uranium environmental availability or in P. magellanicus biomineralization between these two sites. The resolution of this new analytical method raises questions about such data interpretation related to P. magellanicus growth dynamics and physiology or individual scale based environmental measurements.

A current synthesis on the effects of electric and magnetic fields emitted by submarine power cables on invertebrates.

The goal of clean renewable energy production has promoted the large-scale deployment of marine renewable energy devices, and their associated submarine cable network. Power cables produce both electric and magnetic fields that raise environmental concerns as many marine organisms have magneto and electroreception abilities used for vital purposes. Magnetic and electric fields' intensities decrease with distance away from the cable. Accordingly, the benthic and the sedimentary compartments are exposed to the highest field values. Although marine invertebrate species are the major fauna of these potentially exposed areas, they have so far received little attention. We provide extensive background knowledge on natural and anthropogenic marine sources of magnetic and electric fields. We then compile evidence for magneto- and electro-sensitivity in marine invertebrates and further highlight what is currently known about their interactions with artificial sources of magnetic and electric fields. Finally we discuss the main gaps and future challenges that require further investigation.

Compound-specific recording of gadolinium pollution in coastal waters by great scallops.

Gadolinium-based contrast agents (GBCAs), routinely used in magnetic resonance imaging (MRI), end up directly in coastal seawaters where gadolinium concentrations are now increasing. Because many aquatic species could be sensitive to this new pollution, we have evaluated the possibility of using shellfish to assess its importance. Gadolinium excesses recorded by scallop shells collected in Bay of Brest (Brittany, France) for more than 30 years do not reflect the overall consumption in GBCAs, but are largely controlled by one of them, the gadopentetate dimeglumine. Although its use has been greatly reduced in Europe over the last ten years, gadolinium excesses are still measured in shells. Thus, some gadolinium derived from other GBCAs is bioavailable and could have an impact on marine wildlife.