The biology and ecology of the basking shark: A review.

Here we review the literature on the basking shark (Cetorhinus maximus, Gunnerus, 1765), well known as the second largest extant shark (and fish) species globally. Previous reviews were published by Kunzlik in 1988 and Sims in 2008, but in the last 15 years modern electronic and DNA sequencing technologies have resulted in considerable advances in our knowledge of the species' behaviour and ecology. Basking sharks are planktivores and under appropriate conditions spend prolonged periods at the ocean surface feeding on copepod prey that primarily make up their diet, the behaviour that gave rise to their common name. In general, they are migratory and move into higher latitude waters during the summer months, when loose surface-feeding aggregations may form at favoured sites, the best known of which at present occur at hotspots on the west coasts of Britain and Ireland. The species is found circumglobally in temperate waters, but they are also now known on occasion to migrate at depth between northern and southern hemispheres, as well as across oceans within the northern hemisphere. In the past basking shark were more abundant across much of their range, but, consequent on targeted fisheries and in some places intentional eradication, became everywhere scarce, with recent population recovery in the north-east Atlantic being the result of protective measures initiated in the 1990s. Despite their charismatic nature, some of their most fundamental biological processes including copulation, gestation and birth remain largely unknown, due to their migratory and often deep-water lifestyle. In contrast, the deployment of small-scale archival and satellite tags has revealed the details of both broadscale migratory movements and horizontal and vertical foraging behaviours. Recent genetic studies support evidence suggesting a degree of site fidelity in relation to seasonal feeding grounds, which likely explains why in the past local populations have collapsed following periods of intensive fishing. Other recent research using aerial drones and towed cameras has revealed within loose feeding aggregations elements of social behaviour that may have a courtship function as well as enhance feeding efficiency.

Individual residency behaviours and seasonal long-distance movements in acoustically tagged Caribbean reef sharks in the Cayman Islands.

Understanding how reef-associated sharks use coastal waters through their ontogeny is important for their effective conservation and management. This study used the horizontal movements of acoustically tagged Caribbean reef sharks (Carcharhinus perezi) to examine their use of coastal space around the Cayman Islands between 2009 and 2019. A total of 39 (59.1%) tagged sharks (male = 22, female = 17, immature = 18, mature = 21) were detected on the islands wide network of acoustic receivers. The detection data were used to calculate values of Residency Index (RI), Site-Fidelity Index (SFI) and minimum linear displacement (MLD), as well as for network analysis of individual shark movements to test for differences between demographics, seasons, and diel periods. Sharks were detected for up to 1,598 days post-tagging and some individuals showed resident behaviour but the majority of tagged individuals appear to have been one-off or only occasional transient visitors to the area. Generally, individuals showed strong site-fidelity to different areas displaying linear home ranges of < 20 km. The evidence indicates that there was no pattern of diel behaviour. Tagged sharks generally showed increased movements within and between islands during the summer (April-September), which may be related to breeding activity. Some individuals even made occasional excursions across 110 km of open water > 2,000 m deep between Grand Cayman and Little Cayman. One mature female shark showed a displacement of 148.21 km, the greatest distance reported for this species. The data shows that the distances over which some sharks moved, greatly exceeded the extent of any one of the islands' marine protected areas indicating that this species may be more mobile and dispersive than previously thought. This study provides support for the blanket protection to all sharks throughout Cayman waters, which was incorporated within the National Conservation Act in 2015.

Coral bacterial community structure responds to environmental change in a host-specific manner.

The global decline of coral reefs heightens the need to understand how corals respond to changing environmental conditions. Corals are metaorganisms, so-called holobionts, and restructuring of the associated bacterial community has been suggested as a means of holobiont adaptation. However, the potential for restructuring of bacterial communities across coral species in different environments has not been systematically investigated. Here we show that bacterial community structure responds in a coral host-specific manner upon cross-transplantation between reef sites with differing levels of anthropogenic impact. The coral Acropora hemprichii harbors a highly flexible microbiome that differs between each level of anthropogenic impact to which the corals had been transplanted. In contrast, the microbiome of the coral Pocillopora verrucosa remains remarkably stable. Interestingly, upon cross-transplantation to unaffected sites, we find that microbiomes become indistinguishable from back-transplanted controls, suggesting the ability of microbiomes to recover. It remains unclear whether differences to associate with bacteria flexibly reflects different holobiont adaptation mechanisms to respond to environmental change.

Similar bacterial communities on healthy and injured skin of black tip reef sharks.

BACKGROUND: Sharks are in severe global decline due to human exploitation. The additional concern of emerging diseases for this ancient group of fish, however, remains poorly understood. While wild-caught and captive sharks may be susceptible to bacterial and transmissible diseases, recent reports suggest that shark skin may harbor properties that prevent infection, such as a specialized ultrastructure or innate immune properties, possibly related to associated microbial assemblages. To assess whether bacterial community composition differs between visibly healthy and insulted (injured) shark skin, we compared bacterial assemblages of skin covering the gills and the back from 44 wild-caught black-tip reef sharks (Carcharhinus melanopterus) from the Amirante Islands (Seychelles) via 16S rRNA gene amplicon sequencing. RESULTS: Shark skin-associated bacterial communities were diverse (5971 bacterial taxa from 375 families) and dominated by three families of the phylum Proteobacteria typical of marine organisms and environments (Rhodobacteraceae, Alteromonadaceae, Halomonadaceae). Significant differences in bacterial community composition of skin were observed for sharks collected from different sites, but not between healthy or injured skin samples or skin type (gills vs. back). The core microbiome (defined as bacterial taxa present in ≥50% of all samples) consisted of 12 bacterial taxa, which are commonly observed in marine organisms, some of which may be associated with animal host health. CONCLUSION: The conserved bacterial community composition of healthy and injured shark skin samples suggests absence of severe bacterial infections or substantial pathogen propagation upon skin insult. While a mild bacterial infection may have gone undetected, the overall conserved bacterial community implies that bacterial function(s) may be maintained in injured skin. At present, the contribution of bacteria, besides intrinsic animal host factors, to counter skin infection and support rapid wound healing in sharks are unknown. This represents clear knowledge gaps that should be addressed in future work, e.g. by screening for antimicrobial properties of skin-associated bacterial isolates.

Some like it hot: Repeat migration and residency of whale sharks within an extreme natural environment.

The Arabian Gulf is the warmest sea in the world and is host to a globally significant population of the whale shark Rhincodon typus. To investigate regional whale shark behaviour and movements, 59 satellite-linked tags were deployed on whale sharks in the Al Shaheen area off Qatar from 2011-14. Four different models of tag were used throughout the study, each model able to collect differing data or quantities of data. Retention varied from one to 227 days. While all tagged sharks crossed international maritime boundaries, they typically stayed within the Arabian Gulf. Only nine sharks dispersed through the narrow Strait of Hormuz into the Gulf of Oman. Most sharks stayed close to known or suspected feeding aggregation sites over summer months, but dispersed throughout the Arabian Gulf in winter. Sharks rarely ventured into shallow areas (<40 m depth). A single, presumably pregnant female shark was the sole animal to disperse a long distance, crossing five international maritime boundaries in 37 days before the tag detached at a distance of approximately 2644 km from the tagging site, close to the Yemeni-Somali border. No clear space-use differentiation was evident between years, for sharks of different sizes, or between sexes. Whale sharks spent the most time (~66%) in temperatures of 24-30°C and in shallow waters <100 m depth (~60%). Sharks spent relatively more time in cooler (X2 = 121.692; p<0.05) and deeper (X2 = 46.402; p<0.05) water at night. Sharks rarely made dives deeper than 100 m, reflecting the bathymetric constraints of the Gulf environment. Kernel density analysis demonstrated that the tagging site at Al Shaheen was the regional hotspot for these sharks, and revealed a probable secondary aggregation site for whale sharks in nearby Saudi Arabian waters. Analysis of visual re-sightings data of tagged sharks revealed that 58% of tagged individuals were re-sighted back in Al Shaheen over the course of this study, with 40% recorded back at Al Shaheen in the year following their initial identification. Two sharks were confirmed to return to Al Shaheen in each of the five years of study.

Population Structure, Abundance and Movement of Whale Sharks in the Arabian Gulf and the Gulf of Oman.

Data on the occurrence of whale sharks, Rhincodon typus, in the Arabian Gulf and Gulf of Oman were collected by dedicated boat surveys and via a public-sightings scheme during the period from 2011 to 2014. A total of 422 individual whale sharks were photo-identified from the Arabian Gulf and the northern Gulf of Oman during that period. The majority of sharks (81%, n = 341) were encountered at the Al Shaheen area of Qatar, 90 km off the coast, with the Musandam region of Oman a secondary area of interest. At Al Shaheen, there were significantly more male sharks (n = 171) than females (n = 78; X2 = 17.52, P < 0.05). Mean estimated total length (TL) for sharks was 6.90 m ± 1.24 (median = 7 m; n = 296). Males (7.25 m ± 1.34; median = 8 m, n = 171) were larger than females (6.44 m ±1.09; median = 7 m, n = 78; Mann-Whitney U test, p < 0.01). Of the male sharks assessed for maturity 63% were mature (n = 81), with 50% attaining maturity by 7.29 m and 100% by 9.00 m. Two female sharks of >9 m individuals were visually assessed as pregnant. Connectivity among sharks sighted in Qatari, Omani and UAE waters was confirmed by individual spot pattern matches. A total of 13 identified sharks were re-sighted at locations other than that at which they were first sighted, including movements into and out of the Arabian Gulf through the Strait of Hormuz. Maximum likelihood techniques were used to model an estimated combined population for the Arabian Gulf and Gulf of Oman of 2837 sharks ± 1243.91 S.E. (95% C.I. 1720-6295). The Al Shaheen aggregation is thus the first site described as being dominated by mature males while the free-swimming pregnant females are the first reported from the Indian Ocean.

Use of Photo-Identification and Mark-Recapture Methodology to Assess Basking Shark (Cetorhinus maximus) Populations.

Following centuries of exploitation, basking sharks (Cetorhinus maximus) are considered by IUCN as Endangered in the Northeast Atlantic, where they have now been substantially protected for over two decades. However, the present size of this population remains unknown. We investigated the use of photo-identification of individuals' dorsal fins, combined with mark-recapture methodology, to investigate the size of populations of basking shark within the west coast of Scotland. From a total of 921 encounters photographed between 2004 and 2011, 710 sharks were found to be individually identifiable based on dorsal fin damage and natural features. Of these, only 41 individuals were re-sighted, most commonly both within days of, and close to the site of, the initial encounter. A smaller number were re-sighted after longer periods of up to two years. A comparison of the distinguishing features of individuals on first recording and subsequent re-sighting showed that in almost all cases these features remained little changed, suggesting the low re-sighting rate was not due to a loss of distinguishing features. Because of the low number of re-sighting we were not able to produce reliable estimates for the long-term regional population. However, for one 50 km diameter study area between the islands of Mull, Coll and Tiree, we were able to generate closed-population estimates for 6-9 day periods in 2010 of 985 (95% CI = 494-1683), and in 2011 of 201 (95% CI = 143-340). For the same 2011 period an open-population model generated a similar estimate of 213 (95% CI = 111-317). Otherwise the low rate and temporal patterning of re-sightings support the view that such local basking shark populations are temporary, dynamic groupings of individuals drawn from a much larger regional population than previously supposed. The study demonstrated the feasibility and limitations of photo-identification as a non-invasive technique for identifying individual basking sharks.

Coral microbial community dynamics in response to anthropogenic impacts near a major city in the central Red Sea.

Coral-associated bacteria play an increasingly recognized part in coral health. We investigated the effect of local anthropogenic impacts on coral microbial communities on reefs near Jeddah, the largest city on the Saudi Arabian coast of the central Red Sea. We analyzed the bacterial community structure of water and corals (Pocillopora verrucosa and Acropora hemprichii) at sites that were relatively unimpacted, exposed to sedimentation & local sewage, or in the discharge area of municipal wastewaters. Coral microbial communities were significantly different at impacted sites: in both corals the main symbiotic taxon decreased in abundance. In contrast, opportunistic bacterial families, such as e.g. Vibrionaceae and Rhodobacteraceae, were more abundant in corals at impacted sites. In conclusion, microbial community response revealed a measurable footprint of anthropogenic impacts to coral ecosystems close to Jeddah, even though the corals appeared visually healthy.

Sharks and people: insight into the global practices of tourism operators and their attitudes to shark behaviour.

Shark tourism is a popular but controversial activity. We obtained insights into this industry via a global e-mailed questionnaire completed by 45 diving/snorkelling operators who advertised shark experiences (shark operators) and 49 who did not (non-shark operators). 42% of shark operators used an attractant to lure sharks and 93% stated they had a formal code of conduct which 86% enforced "very strictly". While sharks were reported to normally ignore people, 9 operators had experienced troublesome behaviour from them. Whilst our research corroborates previous studies indicating minimal risk to humans from most shark encounters, a precautionary approach to provisioning is required to avoid potential ecological and societal effects of shark tourism. Codes of conduct should always stipulate acceptable diver behaviour and appropriate diver numbers and shark operators should have a moral responsibility to educate their customers about the need for shark conservation.

Whale sharks, Rhincodon typus, aggregate around offshore platforms in Qatari waters of the Arabian Gulf to feed on fish spawn.

Whale sharks, Rhincodon typus, are known to aggregate to feed in a small number of locations in tropical and subtropical waters. Here we document a newly discovered major aggregation site for whale sharks within the Al Shaheen oil field, 90 km off the coast of Qatar in the Arabian Gulf. Whale sharks were observed between April and September, with peak numbers observed between May and August. Density estimates of up to 100 sharks within an area of 1 km(2) were recorded. Sharks ranged between four and eight metres' estimated total length (mean 6.92 ± 1.53 m). Most animals observed were actively feeding on surface zooplankton, consisting primarily of mackerel tuna, Euthynnus affinis, eggs.

Transatlantic migration and deep mid-ocean diving by basking shark.

Despite being the second largest fish, basking sharks (Cetorhinus maximus) have been assumed to remain in discrete populations. Their known distribution encompasses temperate continental shelf areas, yet until now there has been no evidence for migration across oceans or between hemispheres. Here we present results on the tracks and behaviour of two basking sharks tagged off the British Isles, one of which released its tag off Newfoundland, Canada. During the shark's transit of the North Atlantic, she travelled a horizontal distance of 9589 km and reached a record depth of 1264 m. This result provides the first evidence for a link between European and American populations and indicates that basking sharks make use of deep-water habitats beyond the shelf edge.

Visitor perceptions and the shifting social carrying capacity of South Sinai's coral reefs.

To investigate how the perceptions and behaviour of visitors to coral reefs are influenced by their prior experience and knowledge of marine life, a questionnaire-based study was undertaken at sites in the Ras Mohammed National Park and at Sharm El Sheikh, South Sinai, Egypt. It was evident that over the 10-20 years during which these reefs have deteriorated (mainly due to reef-flat trampling), there have been interrelated shifts in the nature of visitors making use of them. First, there has been a shift from experienced divers and snorkellers to inexperienced snorkellers and non-snorkellers with a poorer knowledge of reef biology. Second, there has been a shift in the predominant nationalities of visitors, from German and British, through Italian, to Russian. More recent user groups both stated and showed that they had less experience of snorkelling; they also showed less knowledge of marine life and less interest in learning about it. Visitor perceptions of both the state of the marine life on the reefs and the acceptability of current visitor numbers also varied between groups. More recent visitor groups and visitors with less knowledge were more satisfied with reef health. In general, however, visitor perceptions of reef health did not correlate well with actual reef conditions, probably because more experienced visitors preferred less impacted sites with which they were nevertheless less satisfied than inexperienced visitors at heavily impacted sites. More recent visitor groups were also less bothered by crowding on the shore or in the water. Consequently, the apparent "social carrying capacity" of sites seems to be increasing to a level well above the likely "ecological carrying capacity".