Cetorhinus Research Articles

Bactericidal ability of target acidic phospholipids and phagocytosis of CDC42 GTPase-mediated cytoskeletal rearrangement underlie functional conservation of CXCL12 in vertebrates.

Chemokine CXCL12 plays a crucial role in both direct bactericidal activity and phagocytosis in humans. However, the mechanisms and evolutionary functions of these processes in vertebrates remain largely unknown. In this study, we found that the direct bactericidal activity of CXCL12 is highly conserved across various vertebrate lineages, including Arctic lamprey (Lampetra japonica), Basking shark (Cetorhinus maximus), grass carp (Ctenopharyngodon idella), Western clawed frog (Xenopus tropicalis), Green anole (Anolis carolinensis), chicken (Gallus gallus), and human (Homo sapiens). CXCL12 also has been shown to promote phagocytosis in lower and higher vertebrates. We then employed C. idella CXCL12a (CiCXCL12a) as a model to further investigate its immune functions and underlying mechanisms. CiCXCL12a exerts direct broad-spectrum antibacterial activity by targeting bacterial acidic phospholipids, resulting in bacterial cell membrane perforation, and eventual lysis. Monocytes/macrophages are attracted to the infection sites for phagocytosis through the rapid production of CiCXCL12a during bacterial infection. CiCXCL12a induces CDC42 and CDC42 GTPase activation, which in turn mediates F-actin polymerization and cytoskeletal rearrangement. The interaction between F-actin and Aeromonas hydrophila facilitates bacterial internalization into monocytes/macrophages. Additionally, A. hydrophila is colocalized within early endosomes, late endosomes and lysosomes, ultimately degrading within phagolysosomes. CiCXCL12a also activates PI3K-AKT, JAK-STAT5 and MAPK-ERK signaling pathways. Notably, only the PI3K-AKT signaling pathway inhibits LPS-induced monocyte/macrophage apoptosis. Thus, CiCXCL12a plays key roles in reducing tissue bacterial loads, attenuating organ injury, and decreasing mortality rates. Altogether, our findings elucidate the conserved mechanisms underlying CXCL12-mediated bactericidal activity and phagocytosis, providing novel perspectives into the immune functions of CXCL12 in vertebrates.

Behavioral response of megafauna to boat collision measured via animal-borne camera and IMU

Overlap between marine megafauna and maritime activities is a topic of global concern. Basking sharks (Cetorhinus maximus; CM) are listed as Globally Endangered under the IUCN, though reported sightings appear to be increasing in Ireland. While such trends in the region are welcome, increasing spatiotemporal overlap between CM and numerous water users poses an increased risk of boat strikes to the animals. To demonstrate the risk and impact of boat strikes on marine megafauna, we present camera-enabled animal-borne inertial measurement unit (IMU) data from a non-lethal boat strike on a CM within a proposed National Marine Park in Ireland. We tagged a ~7-m female CM in County Kerry, Ireland, which was struck by a boat ~6 h after tag deployment. Comparison of pre-strike data with 4 h of video and ~7.5 h of IMU data following the boat strike provides critical insight into the animal’s response. While the CM reacted momentarily with an increase in activity and swam to the seafloor, it quickly reduced its overall activity (i.e., overall dynamic body acceleration, tailbeat cycles, tailbeat amplitude, and vertical velocity) for the remainder of the deployment. Notably, the animal also ceased feeding for the duration of the video and headed towards deep offshore waters, which is in stark contrast to the pre-strike period where the animal was consistently observed feeding along the surface in shallow coastal water. This work provides insight into a CM’s response to acute injury and highlights the need for appropriate protections to mitigate risks for marine megafauna.

Assessing the potential of acoustic telemetry to underpin the regional management of basking sharks (Cetorhinus maximus)

Acoustic telemetry can provide valuable space-use data for a range of marine species. Yet the deployment of species-specific arrays over vast areas to gather data on highly migratory vertebrates poses formidable challenges, often rendering it impractical. To address this issue, we pioneered the use of acoustic telemetry on basking sharks (Cetorhinus maximus) to test the feasibility of using broadscale, multi-project acoustic receiver arrays to track the movements of this species of high conservation concern through the coastal waters of Ireland, Northern Ireland, and Scotland. Throughout 2021 and 2022, we tagged 35 basking sharks with acoustic transmitters off the west coast of Ireland; 27 of these were detected by 96 receiver stations throughout the study area (n = 9 arrays) with up to 216 detections of an individual shark (mean = 84, s.d. 65). On average, sharks spent ~ 1 day at each acoustic array, with discrete residency periods of up to nine days. Twenty-one sharks were detected at multiple arrays with evidence of inter-annual site fidelity, with the same individuals returning to the same locations in Ireland and Scotland over 2 years. Eight pairs of sharks were detected within 24 h of each other at consecutive arrays, suggesting some level of social coordination and synchronised movement. These findings demonstrate how multi-project acoustic telemetry can support international, cost-effective monitoring of basking sharks and other highly mobile species. Decision support tools such as these can consolidate cross-border management strategies, but to achieve this goal, collaborative efforts across jurisdictions are necessary to establish the required infrastructure and secure ongoing support.

Novel pre-copulatory behavior in basking sharks observed by drone.

Basking sharks (Cetorhinus maximus) seasonally aggregate in coastal surface waters of the North Atlantic, providing opportunities for visual observation. While putative courtship displays have been observed, actual copulation has not been documented. Here we examine video collected by an unmanned aerial vehicle ("drone") of novel behavioral interactions between basking sharks in Cape Cod Bay, Massachusetts in May 2021. The behaviors, including close following and tight concentric circling, are consistent with pre-copulatory behavior observed in other shark species. These observations provide new insights into the pre-copulatory behavior of basking sharks.

On Sharks Unseen

Basking sharks are the planet’s second-largest fish, and in the summer they feed on plankton in the Sea of the Hebrides, in Scotland. Once hunted for the oil contained in their livers, basking sharks are now a protected species, with tour companies offering the possibility to see and even snorkel with them. There is no guarantee of a sighting, however. This essay takes as its point of departure one such unsuccessful attempt to find basking sharks, undertaken as part of a research trip to learn about the history of shark hunting in the north-east Atlantic. Engaging with literature from multispecies ethnography, the essay considers the implications of treating absences as a condition of research on underwater species. It asks what form the tenets of multispecies ethnography – such as arts of attentiveness, immersion, and sustained participation in the lives of others – can take in oceanic settings. It suggests that direct observation cannot always account for relations with the unseen, and that methodologically and conceptually, the non-encounter offers a way of thinking through the ways that human activity can contribute to the loss of other species.

Allometric Growth of the Enigmatic Deep-Sea Megamouth Shark Megachasma pelagios Taylor, Compagno, and Struhsaker, 1983 (Lamniformes, Megachasmidae)

Megamouth sharks Megachasma pelagios Taylor, Compagno, and Struhsaker, 1983, are a large-bodied, planktivorous, deep-sea species with peculiar morphology. Since their initial description in the late 20th century, many individuals of different sizes have been reported, but few studies examined ontogenetic changes in body shapes. Here, we assess the growth changes in their heads and fins based on length measurements from nine different-sized individuals (177–544 cm in total length). Bivariate analyses showed that the head becomes larger relative to body length with increasing body size (i.e., positive allometry), whereas the relative size of the caudal fin remains constant (i.e., isometric growth). This trend differs from basking sharks and apparently resembles whale sharks and some baleen whales, although they are all large-bodied filter feeders. Given that relative mouth size is linked to feeding modes, our results suggest that megamouth sharks have different feeding modes from ram-feeding basking sharks and may have some similarity with suction-feeding whale sharks and engulfment-feeding baleen whales.

Hydrodynamic analysis of bioinspired vortical cross-step filtration by computational modelling.

Research on the suspension-feeding apparatus of fishes has led recently to the identification of novel filtration mechanisms involving vortices. Structures inside fish mouths form a series of 'backward-facing steps' by protruding medially into the mouth cavity. In paddlefish and basking shark mouths, porous gill rakers lie inside 'slots' between the protruding branchial arches. Vortical flows inside the slots of physical models have been shown to be important for the filtration process, but the complex flow patterns have not been visualised fully. Here we resolve the three-dimensional hydrodynamics by computational fluid dynamics simulation of a simplified mouth cavity including realistic flow dynamics at the porous layer. We developed and validated a modelling protocol in ANSYS Fluent software that combines a porous media model and permeability direction vector mapping. We found that vortex shape and confinement to the medial side of the gill rakers result from flow resistance by the porous gill raker surfaces. Anteriorly directed vortical flow shears the porous layer in the centre of slots. Flow patterns also indicate that slot entrances should remain unblocked, except for the posterior-most slot. This new modelling approach will enable future design exploration of fish-inspired filters.

Regionally endothermic traits in planktivorous basking sharks Cetorhinus maximus

Few fast-swimming apex fishes are classified as ‘regional endotherms’, having evolved a relatively uncommon suite of traits (e.g. elevated body temperatures, centralised red muscle, and thick-walled hearts) thought to facilitate a fast, predatory lifestyle. Unlike those apex predators, Endangered basking sharks Cetorhinus maximus are massive filter-feeding planktivores assumed to have the anatomy and physiology typical of fully ectothermic fishes. We combined dissections of stranded specimens with biologging of free-swimming individuals and found that basking sharks have red muscle located medially at the trunk, almost 50% compact myocardium of the ventricle, and subcutaneous white muscle temperatures consistently 1.0 to 1.5°C above ambient. Collectively, our findings suggest basking sharks are not full ectotherms, instead sharing several traits used to define a regional endotherm, thus deviating from our current understanding of the species and questioning the link between physiology and ecology of regionally endothermic shark species. With successful forecasting of population dynamics and distribution shifts often improved by accurate physiological data, our results may help explain movement patterns of the species, which could ultimately facilitate conservation efforts.

Highly variable basking shark (Cetorhinus maximus) diving behavior in the lower Bay of Fundy, Canada

Diving behavior in basking sharks, the largest obligate ram filter feeding planktivore, is highly dependent on their location. In the Bay of Fundy, where basking sharks congregate in the boreal summer and autumn, the sharks’ copepod prey are located deep in the water column, below 100 m, in dense but scattered patches. We used time-depth recorders to examine how the vertical movements of basking sharks adapt to such a prey field and captured 4,138 hours of diving behavior from 42 sharks in the boreal summer from 2008 to 2020. Using finite mixture models, we split the time series into surface and subsurface movement blocks and used dynamic time-warping to cluster subsurface movements into seven modes based on their shapes and lengths, with mostly V-shaped subsurface movements (85%) and a minority that were U-shaped (14%). Across sharks, five overall strategies of vertical movement behavior were identified. The strategies split broadly by the ratio of V-shaped movements to U-shaped movements in a deployment and whether the majority of subsurface movements were above or below 100 m. A majority of basking sharks (64%) were reverse diel vertical migrators but none altered their time-allocation across tidal periods. During more thermal stratification, sharks dove deeper, longer, and less frequently while during less thermal stratification sharks dove shallower, shorter, and more frequently. Overall, we show that basking sharks exhibit considerable inter- and intra-individual variability in their diving behavior, and therefore presumably also in foraging modes. Some of this variability relates to time of year and tidal phase, unsurprising in this highly tidally-driven system; however, the majority of the variability remains unexplained without more information on the distribution, composition, and abundance of the copepod prey field. The technique presented is extendable to other species and, unlike many dive classification techniques, requires few subjective delineations of diving behavior.

Circles in the sea: annual courtship "torus" behaviour of basking sharks Cetorhinus maximus identified in the eastern North Atlantic Ocean.

Groups of basking sharks engaged in circling behaviour are rarely observed, and their function remains enigmatic in the absence of detailed observations. Here, underwater and aerial video recordings of multiple circling groups of basking sharks during late summer (August and September 2016-2021) in the eastern North Atlantic Ocean showed groups numbering between 6 and 23 non-feeding individuals of both sexes. Sharks swam slowly in a rotating "torus" (diameter range: 17-39 m), with individuals layered vertically from the surface to a maximum depth of 16 m. Within a torus, sharks engaged in close-following, echelon, close-flank approach or parallel-swimming behaviours. Measured shark total body lengths were 5.4-9.5m (mean LT : 7.3m ± 0.9s.d.; median: 7.2m, n= 27), overlapping known lengths of sexually mature males and females. Males possessed large claspers with abrasions that were also observed on female pectoral fins. Female body colouration was paler than that of males, similar to colour changes observed during courtship and mating in other shark species. Individuals associated with most other members rapidly (within minutes), indicating toroidal behaviours facilitate multiple interactions. Sharks interacted through fin-fin and fin-body contacts, rolling to expose the ventral surfaces to following sharks, and breaching behaviour. Toruses formed in late summer when feeding aggregations in zooplankton-rich thermal fronts switched to non-feeding following and circling behaviours. Collectively, the observations explain a courtship function for toruses. This study highlights northeast Atlantic coastal waters as a critical habitat supporting courtship reproductive behaviour of endangered basking sharks, the first such habitat identified for this species globally.

Legacy and emerging contaminants in the endangered filter feeder basking shark Cetorhinus maximus

The status of contamination by chemical pollutants on large filter feeding sharks is still largely unknown. This study investigated for the first time the presence of legacy, emerging contaminants and trace elements in multiple tissues of basking sharks. In general, skin showed higher concentration of legacy and emerging contaminants probably due to pollutants being adsorbed onto the dermal denticles of the skin rather than accumulated in the tissue itself. Contaminants measured in both subcutaneous tissue and muscles appeared to strongly correlate with each other, indicating that the former might be a good proxy of muscle contamination in basking sharks. Considering the migratory nature of this species, longevity and feeding ecology, this species represents the perfect candidate to act as early warning bioindicator of regional contamination. In this context, non-lethal subcutaneous biopsies could allow the early detection of any temporal variation in the bioaccumulation of pollutants in the Mediterranean Sea.

Spatial Distribution, Temporal Changes, and Knowledge Gaps in Basking Shark (Cetorhinus maximus) Sightings in the California Current Ecosystem

Among the largest fish species, the basking shark (Cetorhinus maximus) is found circumglobally in temperate and tropical waters. Though historical documents have recorded their presence in the California Current Ecosystem (CCE), basking sharks are now only rarely observed in this part of their range. We compiled recent and historical data from systematic surveys (1962–1997) and other sources (1973–2018) to (i) examine temporal patterns of basking shark sightings in the CCE, and (ii) determine the spatial, temporal, and environmental drivers that have affected basking shark presence and distribution here for the last 50 years. We first calculated variation in basking shark sightings and school size over time. We then generated species distribution models using the systematic survey data and evaluated the performance of these models against the more recent non-systematic sightings data. The sightings records indicated that the number of shark sightings was variable across years, but the number and probability of sightings declined in the mid-1980s. The systematic survey data showed up to nearly 4,000 sharks sighted per year until the 1990s, after which there were no sightings reported. In parallel, there was more than a 50% decline in school size from the 1960s to the 1980s (57.2 to 24.0 individuals per group). During the subsequent decades in the non-systematic data (>1990), less than 60 sharks were sighted per year. There were no schools larger than 10 reported, and the mean school size in the last decade (2010s) was 3.53 individuals per group. Low sea surface temperature and high chlorophyllaconcentration increased sightings probability, and prevailing climatic oscillations (El Nino-Southern Oscillation index, North Pacific Gyre Oscillation, Pacific Decadal Oscillation) were also correlated with basking shark presence. Lastly, we observed a significant shift in the seasonality of sightings, from the fall and spring during the systematic survey period to the summer months after the 2000s. We conclude by offering suggestions for future research and conservation efforts; specifically, coordinating the documentation of fisheries mortalities and sightings throughout the Pacific basin would facilitate more robust population estimates and identify sources of mortality. Additionally, monitoring shark fin markets and developing region-specific genetic markers would help ensure that convention on international trade in endangered species (CITES) regulations are being followed.

Basking shark tourism in Donegal, Ireland – a case‐study of public interest and support for shark conservation

Abstract Basking sharks (Cetorhinus maximus) visit the coastal waters of Ireland annually, and while protected in Northern Ireland, they are not protected by domestic legislation in Ireland. In Ireland, basking shark conservation relies heavily on publicly reported sightings to assess population size and migration patterns. While basking shark tourism is popular in nearby Scotland, Irish tourism materials rarely feature basking sharks. In order to determine public awareness about basking sharks, public support for basking shark conservation and interest in shark tourism, 173 residents and visitors in Buncrana, Ireland were surveyed in July 2018. The results indicated that public knowledge of basking sharks is low, whilst support for basking shark conservation and interest in tourism are high. Despite the fact that conservation groups rely on publicly reported sightings as a cost‐effective research tool that is important for conservation policy, only 7% of survey respondents were aware that they could report a basking shark sighting to local research organizations (Irish Whale and Dolphin Group or Irish Basking Shark Group). Individuals who support conservation were significantly more likely to believe that others would be willing to pay to view basking sharks, indicating a link between tourism potential and support for conservation. This study showed that there may be an untapped tourism market for basking shark viewing in Donegal as well as public support for shark conservation.

Cool runnings: behavioural plasticity and the realised thermal niche of basking sharks

Long-distance migrations by marine vertebrates are often triggered by pronounced environmental cues. For the endangered basking shark (Cetorhinus maximus), seasonal changes in water temperature are frequently proposed as a cue for aggregation within (and dispersal from) coastal hotspots. The inference is that such movements reflect year-round occupancy within a given thermal ‘envelope’. However, the marked variance in timing, direction and depth of dispersal movements hint at a more nuanced explanation for basking sharks. Here, using data from pop-off archival transmitters deployed on individuals in Irish waters, we explored whether autumnal decreases in water temperature triggered departure from coastal habitats and how depth and location shaped the sharks’ realised thermal environment over time. Temperature was not an apparent driver of dispersal from coastal seas, and variance in daily temperature ranges reflected occupancy of different habitats; coastal mixed/stratified and offshore subtropical/tropical waters. Furthermore, individuals that moved offshore and into more southern latitudes off Africa, exhibited a distinct daily cycle of deep dives (00:00–12:00, 200 m–700 m; 12:00–00:00, 0–300 m), experiencing a more extreme range of temperatures (6.8–27.4 °C), including cooler minimum temperatures, than those remaining in European coastal habitat (9.2–17.6 °C). Collectively, these findings challenge the supposition that temperature serves as a universal driver of seasonal dispersal from coastal seas and prompts further studies of deep-water forays in offshore areas.

The ‘Stronsay Beast’: testimony, evidence and authority in early nineteenth-century natural history

When an unknown sea creature was washed ashore on the Orkney Islands in September 1808, the Edinburgh anatomist John Barclay declared that this was the first solid scientific evidence for the existence of the ‘great sea snake’. The testimony of witnesses along with some of its preserved body parts were examined by both the Wernerian Natural History Society in Edinburgh and the surgeon and anatomist Everard Home in London. Contradicting Barclay's opinion, Home identified the creature as a decomposing basking shark. While Barclay took the testimony of the local witnesses largely on trust and accepted their interpretation of the Beast, Home discounted it and instead asserted his own expert authority to correctly interpret the evidence. Both made use of the preserved physical remains of parts of the creature in strikingly different ways: Barclay to support the accounts of the witnesses, Home to undermine them. The debate between the two anatomists has much to tell us about the uses of evidence and testimony in early nineteenth-century natural history, but also has broader resonances for the roles of evidence and authority in science that still remain relevant today.

Record of a juvenile basking shark, Cetorhinus maximus (Lamniformes: Cetorhinidae), from Sagami Bay, central Japan, with a review of worldwide records of juveniles of the species

Record of a juvenile basking shark, Cetorhinus maximus (Lamniformes: Cetorhinidae), from Sagami Bay, central Japan, with a review of worldwide records of juveniles of the species

Megalodon

Megalodon

Greener Synthesis of Pristane by Flow Dehydrative Hydrogenation of Allylic Alcohol Using a Packed-Bed Reactor Charged by Pd/C as a Single Catalyst.

Our previous work established a continuous-flow synthesis of pristane, which is a saturated branched alkane obtained from a Basking Shark. The dehydration of an allylic alcohol that is the key to a tetraene was carried out using a packed-bed reactor charged by an acid–silica catalyst (HO-SAS) and flow hydrogenation using molecular hydrogen via a Pd/C catalyst followed. The present work relies on the additional propensity of Pd/C to serve as an acid catalyst, which allows us to perform a flow synthesis of pristane from the aforementioned key allylic alcohol in the presence of molecular hydrogen using Pd/C as a single catalyst, which is applied to both dehydration and hydrogenation. The present one-column-two-reaction-flow system could eliminate the use of an acid catalyst such as HO-SAS and lead to a significant simplification of the production process.

A closer look at the bycatch of medium‐sized and large sharks in the northern Catalan coast (north‐western Mediterranean Sea): Evidence of an ongoing decline?

Abstract Historically, the Mediterranean Sea supported a rich shark fauna. Presently, however, populations of most shark species have significantly declined, largely due to intense fishing pressure. Interviews with crew members of bottom trawlers, drifting longliners and bottom longliners operating off the Costa Brava (Catalonia, NE Spain) were conducted between October 2016 and July 2017 in order to gather information on the current bycatch rate of several shark species. Interviews covered 41.2% of the fleet and respondents were asked for the bycatch of selected shark species—Alopias vulpinus, Cetorhinus maximus, Galeorhinus galeus, Hexanchus griseus, Isurus oxyrinchus, Mustelus spp., Prionace glauca, and Squalus acanthias—in two distinct time periods. Bottom trawlers captured the highest diversity of species (eight) followed by bottom longliners (seven), and drifting longliners (three). Most respondents (89.7%) declared having captured at least one shark from 2006 to 2016 but only 56.4% declared having captured at least one shark from 2016 to 2017. From 2016 to 2017, the whole fleet captured 89 specimens of H. griseus (95% confidence interval (CI) = 145, 34), 14 of G. galeus (95% CI = 30, 0), 3 of A. vulpinus (95% CI = 8, 0), 3 of I. oxyrinchus (95% CI = 8, 0), 3 of C. maximus (95% CI = 6, 0), and no Mustelus spp. The total bycatch of P. glauca and S. acanthias was uncertain due to extremely loose confidence intervals. A significant decline was perceived by fishermen in the bycatch of C. maximus and S. acanthias, whereas the bycatch of H. griseus was considered to have remained stable. This study suggests a dramatic reduction in the abundance of most of the medium‐sized and large sharks of the Costa Brava and the likely disappearance of Mustelus spp. from the area. Only H. griseus, S. acanthias, and P. glauca are still being bycaught frequently.

Drivers of Spatial Distributions of Basking Shark (Cetorhinus maximus) in the Southwest Pacific

Basking sharks (Cetorhinus maximus) were widely reported throughout New Zealand waters. Once commonly observed, and sometimes in large numbers, basking sharks are now infrequently reported. Basking shark observations are known to be highly variable across years, and their distribution and occurrence have been shown to be influenced by environmental predictors such as thermal fronts, chl-a concentration, and the abundance of prey (zooplankton). Little is known of basking sharks in the South Pacific and more information on distribution, habitat use, and migratory patterns is required to better understand the species’ regional ecology. Here, we used bootstrapped Habitat Suitability Models [HSM, ensembled from Boosted Regression Tree (BRT) and Random Forest (RF) models] to determine the drivers of basking shark distribution, predict habitat suitability and estimated uncertainty in the South Pacific for the first time. High−resolution environmental (1 km2 grid resolution) and biotic data, including inferred prey species, and all available basking shark records across New Zealand’s Exclusive Economic Zone (EEZ) were included in the ensemble HSMs. The most influential driver of modeled basking shark distribution was vertical flux of particulate organic matter at the seabed, which may indicate higher levels of primary production in the surface ocean and higher prey density in the mesopelagic zone and at the seafloor. The BRT and RF models had good predictive power (AUC and TSS > 0.7) and both models performed similarly with low variability in the model fit metrics. Areas of high basking shark habitat suitability included the east and west coasts of the South Island, Puysegur Ridge, and Auckland Island slope. The outputs produced here could be incorporated into future management framework for assessing threat and conservation needs (e.g., spatially explicit risk assessment) for this regionally protected species, as well as providing guidance for future research efforts (e.g., areas of interest for sampling).