What acoustic telemetry can and cannot tell us about fish biology.

Acoustic telemetry (AT) has become ubiquitous in aquatic monitoring and fish biology, conservation, and management. Since the early use of active ultrasonic tracking that required researchers to follow at a distance their species of interest, the field has diversified considerably, with exciting advances in both hydrophone and transmitter technology. Once a highly specialized methodology, however, AT is fast becoming a generalist tool for those wishing to study or conserve fishes, leading to diversifying application by non-specialists. With this transition in mind, we evaluate exactly what AT has become useful for, discussing how the technological and analytical advances around AT can address important questions within fish biology. In doing so, we highlight the key ecological and applied research areas where AT continues to reveal crucial new insights and, in particular, when combined with complimentary research approaches. We provide a comprehensive breakdown of the state of the art for applications of AT, discussing the ongoing challenges, where its strengths lie, and how future developments may revolutionize fisheries management, behavioral ecology and species protection. Through selected papers we illustrate specific applications across the broad spectrum of fish biology. By bringing together the recent and future developments in this field under categories designed to broadly capture many aspects of fish biology, we hope to offer a useful guide for the non-specialist practitioner as they attempt to navigate the dizzying array of considerations and ongoing developments within this diverse toolkit.

From sneaker to bourgeois male: Long-term observations on recognized tompot blenny Parablennius gattorugine individuals reveal new information on their biology.

The behaviour of the tompot blenny Parablennius gattorugine was studied by underwater observation and photography on the south-west coast of the UK over an 11 year period, with >50 individuals identified at two locations by their distinctive skin markings. Observations showed males engaging in rapid swimming behaviour when approached by a female, and younger males using alternative 'sneaker' tactics before becoming 'bourgeois' territory-holders. The noninvasive recognition technique also facilitated detailed observations on territory retention (up to 7 years) and agonistic behaviours among both males and females.

First record of partial albinism in the critically endangered Angelshark (Squatina squatina) (Linnaeus, 1758).

We report the first case of partial albinism in the Critically Endangered angelshark, Squatina squatina. The encounter with this specimen took place while SCUBA diving on the beach of Tufia, located on the east coast of the island of Gran Canaria on 2 April 2021. This is also the first confirmed finding of an albino elasmobranch specimen in the Canary Island archipelago.

The movement ecology of fishes.

Movement of fishes in the aquatic realm is fundamental to their ecology and survival. Movement can be driven by a variety of biological, physiological and environmental factors occurring across all spatial and temporal scales. The intrinsic capacity of movement to impact fish individually (e.g., foraging) with potential knock-on effects throughout the ecosystem (e.g., food web dynamics) has garnered considerable interest in the field of movement ecology. The advancement of technology in recent decades, in combination with ever-growing threats to freshwater and marine systems, has further spurred empirical research and theoretical considerations. Given the rapid expansion within the field of movement ecology and its significant role in informing management and conservation efforts, a contemporary and multidisciplinary review about the various components influencing movement is outstanding. Using an established conceptual framework for movement ecology as a guide (i.e., Nathan et al., 2008: 19052), we synthesized the environmental and individual factors that affect the movement of fishes. Specifically, internal (e.g., energy acquisition, endocrinology, and homeostasis) and external (biotic and abiotic) environmental elements are discussed, as well as the different processes that influence individual-level (or population) decisions, such as navigation cues, motion capacity, propagation characteristics and group behaviours. In addition to environmental drivers and individual movement factors, we also explored how associated strategies help survival by optimizing physiological and other biological states. Next, we identified how movement ecology is increasingly being incorporated into management and conservation by highlighting the inherent benefits that spatio-temporal fish behaviour imbues into policy, regulatory, and remediation planning. Finally, we considered the future of movement ecology by evaluating ongoing technological innovations and both the challenges and opportunities that these advancements create for scientists and managers. As aquatic ecosystems continue to face alarming climate (and other human-driven) issues that impact animal movements, the comprehensive and multidisciplinary assessment of movement ecology will be instrumental in developing plans to guide research and promote sustainability measures for aquatic resources.

Multiyear social stability and social information use in reef sharks with diel fission-fusion dynamics.

Animals across vertebrate taxa form social communities and often exist as fission-fusion groups. Central place foragers (CPF) may form groups from which they will predictably disperse to forage, either individually or in smaller groups, before returning to fuse with the larger group. However, the function and stability of social associations in predatory fish acting as CPFs is unknown, as individuals do not need to return to a shelter yet show fidelity to core areas. Using dynamic social networks generated from acoustic tracking data, we document spatially structured sociality in CPF grey reef sharks at a Pacific Ocean atoll. We show that sharks form stable social groups over multiyear periods, with some dyadic associations consistent for up to 4 years. Groups primarily formed during the day, increasing in size throughout the morning before sharks dispersed from the reef at night. Our simulations suggest that multiple individuals sharing a central place and using social information while foraging (i.e. local enhancement) will outperform non-CPF social foragers. We show multiyear social stability in sharks and suggest that social foraging with information transfer could provide a generalizable mechanism for the emergence of sociality with group central place foraging.

Mobile marine predators: an understudied source of nutrients to coral reefs in an unfished atoll.

Animal movements can facilitate important ecological processes, and wide-ranging marine predators, such as sharks, potentially contribute significantly towards nutrient transfer between habitats. We applied network theory to 4 years of acoustic telemetry data for grey reef sharks (Carcharhinus amblyrhynchos) at Palmyra, an unfished atoll, to assess their potential role in nutrient dynamics throughout this remote ecosystem. We evaluated the dynamics of habitat connectivity and used network metrics to quantify shark-mediated nutrient distribution. Predator movements were consistent within year, but differed between years and by sex. Females used higher numbers of routes throughout the system, distributing nutrients over a larger proportion of the atoll. Extrapolations of tagged sharks to the population level suggest that prey consumption and subsequent egestion leads to the heterogeneous deposition of 94.5 kg d-1 of nitrogen around the atoll, with approximately 86% of this probably derived from pelagic resources. These results suggest that sharks may contribute substantially to nutrient transfer from offshore waters to near-shore reefs, subsidies that are important for coral reef health.

Is the scaling of swim speed in sharks driven by metabolism?

The movement rates of sharks are intrinsically linked to foraging ecology, predator-prey dynamics and wider ecosystem functioning in marine systems. During ram ventilation, however, shark movement rates are linked not only to ecological parameters, but also to physiology, as minimum speeds are required to provide sufficient water flow across the gills to maintain metabolism. We develop a geometric model predicting a positive scaling relationship between swim speeds in relation to body size and ultimately shark metabolism, taking into account estimates for the scaling of gill dimensions. Empirical data from 64 studies (26 species) were compiled to test our model while controlling for the influence of phylogenetic similarity between related species. Our model predictions were found to closely resemble the observed relationships from tracked sharks, providing a means to infer mobility in particularly intractable species.

Environmental stress reduces shark residency to coral reefs.

Coral reef ecosystems are highly threatened and can be extremely sensitive to the effects of climate change. Multiple shark species rely on coral reefs as important habitat and, as such, play a number of significant ecological roles in these ecosystems. How environmental stress impacts routine, site-attached reef shark behavior, remains relatively unexplored. Here, we combine 8 years of acoustic tracking data (2013-2020) from grey reef sharks resident to the remote coral reefs of the Chagos Archipelago in the Central Indian Ocean, with a satellite-based index of coral reef environmental stress exposure. We show that on average across the region, increased stress on the reefs significantly reduces grey reef shark residency, promoting more diffuse space use and increasing time away from shallow forereefs. Importantly, this impact has a lagged effect for up to 16 months. This may have important physiological and conservation consequences for reef sharks, as well as broader implications for reef ecosystem functioning. As climate change is predicted to increase environmental stress on coral reef ecosystems, understanding how site-attached predators respond to stress will be crucial for forecasting the functional significance of altering predator behavior and the potential impacts on conservation for both reef sharks and coral reefs themselves.

Anguillid eels.

Anguillid eels have fascinated humans for centuries, but our knowledge of these mysterious fish is still scant. There are 19 species or subspecies in the genus Anguilla, which are found globally, except in the eastern Pacific and southern Atlantic. Their common label 'freshwater eels' is a misnomer - all anguillids are facultatively catadromous, born in marine environments, developing in continental waters, with a proportion never entering freshwater at all. Anguillid eels have several life history traits that have allowed them to exploit a broad range of habitats. As such, anguillid eels play an important ecological role in both marine and freshwater environments as well as being commercially valuable. Because of this, anguillid eels are under threat from multiple stressors, such as barriers to migration, pollution, parasites, disease, climate change and unsustainable exploitation. Six species are listed as Threatened in the Red List of Threatened Species, and four are listed as Data Deficient. Strengthening conservation and management of these species is essential, and further research provides an exciting opportunity to develop a greater understanding of this mysterious clade of fish.

Tail shape and the swimming speed of sharks.

Trait-based ecology is a rapidly growing approach for developing insights and predictions for data-poor species. Caudal tail fin shape has the potential to reveal much about the energetics, activity and ecology of fishes and can be rapidly measured from field guides, which is particularly helpful for data-sparse species. One outstanding question is whether swimming speed in sharks is related to two morphological traits: caudal fin aspect ratio (CFAR, height2/tail area) and caudal lobe asymmetry ratio (CLAR). We derived both metrics from the species drawings in Sharks of the world (Ebert et al. 2013 Sharks of the world: a fully illustrated guide) and related fin shape to two published datasets of (1) instantaneous swimming speeds (Jacoby et al. 2015 Biol. Lett. 11, 20150781 (doi:10.1098/rsbl.2015.0781)) and (2) cruising speeds (Harding et al. 2021 Funct. Ecol. 35, 1951-1959 (doi:10.1111/1365-2435.13869)) for 28 total unique shark species. Both estimates of swimming speed were positively related to CFAR (and weakly negatively to CLAR). Hence, shark species with larger CFAR and more symmetric tails (low CLAR) tended to be faster-moving and have higher average speeds. This relationship demonstrates the opportunity to use tail shape as an easily measured trait to index shark swimming speed to broader trait-based analyses of ecological function and extinction risk.

First analysis of multiple paternity in an oviparous shark, the small-spotted catshark (Scyliorhinus canicula L.).

Multiple paternity (MP) has been demonstrated in a variety of sharks, although its prevalence and the number of sires per litter vary considerably among species. To date, such analyses have focused on viviparous species that possess only part of the wide spectrum of reproductive strategies developed in elasmobranchs. We analyzed MP in an oviparous species, the small-spotted catshark (Scyliorhinus canicula). In total, 150 neonates originating from 13 different mothers were genotyped using 12 microsatellite loci. MP was commonplace, with progeny from 92% of females sired by multiple males. This result is consistent with the reproductive biology of the species, particularly its protracted breeding season and potential for long-term sperm storage. The significance of these findings is discussed in light of small-spotted catshark behavior, which suggests that the cost of avoiding mating attempts initiated by males may be high and is therefore supportive of convenience polyandry as an explanation for MP. Eggs were followed from the time they were laid to when they hatched, offering a rare opportunity to investigate juvenile development in more detail.

Social networks and the conservation of fish.

Despite our critical dependence on aquatic wildlife, we lack a complete understanding of the drivers of population stability and structure for most fish species. Social network analysis has been increasingly used to investigate animal societies as it explicitly links individual decision-making to population-level processes and demography. While the study of social structure is of great ecological interest, it is also potentially important for species of economic value or of conservation concern. To date however, there has been little focus on how social processes are likely to influence the conservation of fish populations. Here we identify applications for how a social network approach can help address broad fish conservation themes such as population structure, biological invasions or fisheries management. We discuss the burgeoning opportunities offered and challenges still faced by current technologies to integrate social network approaches within fish conservation.

Diving into the vertical dimension of elasmobranch movement ecology.

Knowledge of the three-dimensional movement patterns of elasmobranchs is vital to understand their ecological roles and exposure to anthropogenic pressures. To date, comparative studies among species at global scales have mostly focused on horizontal movements. Our study addresses the knowledge gap of vertical movements by compiling the first global synthesis of vertical habitat use by elasmobranchs from data obtained by deployment of 989 biotelemetry tags on 38 elasmobranch species. Elasmobranchs displayed high intra- and interspecific variability in vertical movement patterns. Substantial vertical overlap was observed for many epipelagic elasmobranchs, indicating an increased likelihood to display spatial overlap, biologically interact, and share similar risk to anthropogenic threats that vary on a vertical gradient. We highlight the critical next steps toward incorporating vertical movement into global management and monitoring strategies for elasmobranchs, emphasizing the need to address geographic and taxonomic biases in deployments and to concurrently consider both horizontal and vertical movements.

Analysing detection gaps in acoustic telemetry data to infer differential movement patterns in fish.

A wide array of technologies are available for gaining insight into the movement of wild aquatic animals. Although acoustic telemetry can lack the fine-scale spatial resolution of some satellite tracking technologies, the substantially longer battery life can yield important long-term data on individual behavior and movement for low per-unit cost. Typically, however, receiver arrays are designed to maximize spatial coverage at the cost of positional accuracy leading to potentially longer detection gaps as individuals move out of range between monitored locations. This is particularly true when these technologies are deployed to monitor species in hard-to-access locations.Here, we develop a novel approach to analyzing acoustic telemetry data, using the timing and duration of gaps between animal detections to infer different behaviors. Using the durations between detections at the same and different receiver locations (i.e., detection gaps), we classify behaviors into "restricted" or potential wider "out-of-range" movements synonymous with longer distance dispersal. We apply this method to investigate spatial and temporal segregation of inferred movement patterns in two sympatric species of reef shark within a large, remote, marine protected area (MPA). Response variables were generated using network analysis, and drivers of these movements were identified using generalized linear mixed models and multimodel inference.Species, diel period, and season were significant predictors of "out-of-range" movements. Silvertip sharks were overall more likely to undertake "out-of-range" movements, compared with gray reef sharks, indicating spatial segregation, and corroborating previous stable isotope work between these two species. High individual variability in "out-of-range" movements in both species was also identified.We present a novel gap analysis of telemetry data to help infer differential movement and space use patterns where acoustic coverage is imperfect and other tracking methods are impractical at scale. In remote locations, inference may be the best available tool and this approach shows that acoustic telemetry gap analysis can be used for comparative studies in fish ecology, or combined with other research techniques to better understand functional mechanisms driving behavior.

Sea surface temperature dictates movement and habitat connectivity of Atlantic cod in a coastal fjord system.

While movements of organisms have been studied across a myriad of environments, information is often lacking regarding spatio-seasonal patterning in complex temperate coastal systems. Highly mobile fish form an integral part of marine food webs providing linkages within and among habitats, between patches of habitats, and at different life stages. We investigated how movement, activity, and connectivity patterns of Atlantic cod (Gadus morhua) are influenced by dynamic environmental conditions. Movement patterns of 39 juvenile and subadult Atlantic cod were assessed in two coastal sites in the Swedish Skagerrak for 5 months. We used passive acoustic telemetry and network analysis to assess seasonal and spatial movement patterns of cod and their relationships to different environmental factors, using statistical correlations, analysis of recurrent spatial motifs, and generalized linear mixed models. Temperature, in combination with physical barriers, precludes significant connectivity (complex motifs) within the system. Sea surface temperature had a strong influence on connectivity (node strength, degree, and motif frequency), where changes from warmer summer waters to colder winter waters significantly reduced movement activity of fish. As the seasons changed, movement of fish gradually decreased from large-scale (km) linkages in the summer to more localized movement patterns in the winter (limited to 100s m). Certain localized areas, however, were identified as important for connectivity throughout the whole study period, likely due to these multiple-habitat areas fulfilling functions required for foraging and shelter. This study provides new knowledge regarding inshore movement dynamics of juvenile and subadult Atlantic cod that use complex, coastal fjord systems. The findings show that connectivity, seasonal patterns in particular, should be carefully considered when selecting conservation areas to promote marine stewardship.