Exceptionally preserved shark fossils from Mexico elucidate the long-standing enigma of the Cretaceous elasmobranch Ptychodus.

The fossil fish Ptychodus Agassiz, 1834, characterized by a highly distinctive grinding dentition and an estimated gigantic body size (up to around 10 m), has remained one of the most enigmatic extinct elasmobranchs (i.e. sharks, skates and rays) for nearly two centuries. This widespread Cretaceous taxon is common in Albian to Campanian deposits from almost all continents. However, specimens mostly consist of isolated teeth or more or less complete dentitions, whereas cranial and post-cranial skeletal elements are very rare. Here we describe newly discovered material from the early Late Cretaceous of Mexico, including complete articulated specimens with preserved body outline, which reveals crucial information on the anatomy and systematic position of Ptychodus. Our phylogenetic and ecomorphological analyses indicate that ptychodontids were high-speed (tachypelagic) durophagous lamniforms (mackerel sharks), which occupied a specialized predatory niche previously unknown in fossil and extant elasmobranchs. Our results support the view that lamniforms were ecomorphologically highly diverse and represented the dominant group of sharks in Cretaceous marine ecosystems. Ptychodus may have fed predominantly on nektonic hard-shelled prey items such as ammonites and sea turtles rather than on benthic invertebrates, and its extinction during the Campanian, well before the end-Cretaceous crisis, might have been related to competition with emerging blunt-toothed globidensine and prognathodontine mosasaurs.

Warming waters lead to increased habitat suitability for juvenile bull sharks (Carcharhinus leucas).

Coastal ecosystems are highly vulnerable to the impacts of climate change and other stressors, including urbanization and overfishing. Consequently, distributions of coastal fish have begun to change, particularly in response to increasing temperatures linked to climate change. However, few studies have evaluated how natural and anthropogenic disturbances can alter species distributions in conjunction with geophysical habitat alterations, such as changes to land use and land cover (LU/LC). Here, we examine the spatiotemporal changes in the distribution of juvenile bull sharks (Carcharhinus leucas) using a multi-decadal fishery-independent survey of coastal Alabama. Using a boosted regression tree (BRT) modeling framework, we assess the covariance of environmental conditions (sea surface temperature, depth, salinity, dissolved oxygen, riverine discharge, Chl-a) as well as historic changes to LU/LC to the distribution of bull sharks. Species distribution models resultant from BRTs for early (2003-2005) and recent (2018-2020) monitoring periods indicated a mean increase in habitat suitability (i.e., probability of capture) for juvenile bull sharks from 0.028 to 0.082, concomitant with substantial increases in mean annual temperature (0.058°C/yr), Chl-a (2.32 mg/m3), and urbanization (increased LU/LC) since 2000. These results align with observed five-fold increases in the relative abundance of juvenile bull sharks across the study period and demonstrate the impacts of changing environmental conditions on their distribution and relative abundance. As climate change persists, coastal communities will continue to change, altering the structure of ecological communities and the success of nearshore fisheries.

Bacterial abscesses in whitespotted bamboo sharks (Chiloscyllium plagiosum) in managed care.

Sixteen whitespotted bamboo sharks (Chiloscyllium plagiosum) with histologically similar bacterial abscesses were submitted to Northwest ZooPath from nine zoo and aquarium institutions over a 17-year period. These abscesses were characterized by inflammatory cell infiltrates and necrosis with intralesional small, Gram-positive, acid-fast negative, cocci bacteria. The clinical presentation, histologic findings, and culture results indicate that Enterococcus faecalis is a relatively common cause of these lesions in whitespotted bamboo sharks. This organism also provides a treatment challenge due to its inherent antibiotic-resistant properties and ability to form biofilms, confounding the host's immune response. Enterococcus faecalis represents an important cause for abscess formation and cellulitis in captive whitespotted bamboo sharks.

Centralized red muscle in Odontaspis ferox and the prevalence of regional endothermy in sharks.

The order Lamniformes contains charismatic species such as the white shark Carcharodon carcharias and extinct megatooth shark Otodus megalodon, and is of particular interest given their influence on marine ecosystems, and because some members exhibit regional endothermy. However, there remains significant debate surrounding the prevalence and evolutionary origin of regional endothermy in the order, and therefore the development of phenomena such as gigantism and filter-feeding in sharks generally. Here we show a basal lamniform shark, the smalltooth sand tiger shark Odontaspis ferox, has centralized skeletal red muscle and a thick compact-walled ventricle; anatomical features generally consistent with regionally endothermy. This result, together with the recent discovery of probable red muscle endothermy in filter feeding basking sharks Cetorhinus maximus, suggests that this thermophysiology is more prevalent in the Lamniformes than previously thought, which in turn has implications for understanding the evolution of regional endothermy, gigantism, and extinction risk of warm-bodied shark species both past and present.

DEXMEDETOMIDINE AND MIDAZOLAM INTRAMUSCULAR SEDATION IN BROWNBANDED BAMBOO SHARKS (CHILOSCYLLIUM PUNCTATUM).

Bamboo sharks are some of the most common elasmobranch species in zoos and aquaria and are frequently sedated for medical exams, treatments, and research. This study investigated the use of an IM sedation protocol of a single dose of dexmedetomidine (0.05 mg/kg) and midazolam (2.0 mg/kg) in brownbanded bamboo sharks (Chiloscyllium punctatum). Sharks were serially monitored every 5 min for heart rate, branchial beats, righting reflex, coelomic response, cloacal response, pelvic fin reflex, response to noxious stimulus, voluntary movement, and ability to swim. This sedation dose was effective at rapidly and significantly decreasing responses to tactile and noxious stimuli with minimal respiratory depression and was quickly reversible with atipamezole (0.5 mg/kg) and flumazenil (0.05 mg/kg). Sedated sharks developed a mild metabolic acidosis evidenced by a significant increase in lactic acid (mean < 0.37 mmol/L presedation, 4.2 mmol/L after reversal) and decrease in blood pH (mean 7.464 presedation, 7.277 after reversal); however, clinical intervention was not required. This protocol should be further investigated in different elasmobranch species but is promising for providing sedation for noninvasive procedures in brownbanded bamboo sharks.

Associations between total mercury, trace minerals, and blood health markers in Northwest Atlantic white sharks (Carcharodon carcharias).

The ecology and life-histories of white sharks make this species susceptible to mercury bioaccumulation; however, the health consequences of mercury exposure are understudied. We measured muscle and plasma total mercury (THg), health markers, and trace minerals in Northwest Atlantic white sharks. THg in muscle tissue averaged 10.0 mg/kg dry weight, while THg in blood plasma averaged 533 µg/L. THg levels in plasma and muscle were positively correlated with shark precaudal length (153-419 cm), and THg was bioaccumulated proportionally in muscle and plasma. Nine sharks had selenium:mercury molar ratios in blood plasma >1.0, indicating that for certain individuals the potential protective effects of the trace mineral were diminished, whereas excess selenium may have protected other individuals. No relationships between plasma THg and any trace minerals or health markers were identified. Thus, we found no evidence of negative effects of Hg bioaccumulation, even in sharks with very high THg.

Feeding ecology and reproductive biology of small coastal sharks in Malaysian waters.

Small coastal demersal sharks form a major proportion of the sharks landed in Malaysia. However, little is known about their feeding ecology and reproduction. This study sought to elucidate the dietary patterns, role of ontogeny in prey consumption, and reproductive biology of four dominant small demersal shark species in Malaysian waters: the Hasselt's bamboo shark, Chiloscyllium hasseltii; brownbanded bamboo shark, C. punctatum; spadenose shark, Scoliodon laticaudus; and Pacific spadenose shark, S. macrorhynchos. Dietary analyses revealed a high overlap in prey taxa consumed; clear resource partitioning among co-occurring species based on the percentage Prey-specific Index of Relative Importance (%PSIRI), with higher fish %PSIRI for Chiloscyllium hasseltii, higher cephalopod %PSIRI for C. punctatum, and higher crustacean %PSIRI for both Scoliodon species; and an ontogenetic diet shift, seen through changes in prey size. Based on the examination of reproductive organs, the results showed larger sizes at maturity for males compared to females for all four species; no obvious reproductive cycles, based on hepatosomatic and gonadosomatic indices for all species; female bias in the sex ratio of the embryos of Scoliodon species; and increased reproductive output (number of eggs or embryos and size of eggs) with larger female size for C. hasseltii and Scoliodon species. The partitioning of food resources minimizes direct competition for food and supports coexistence within shared coastal habitats. The reproductive strategies of these small coastal sharks appear to be favorable for supporting short-term population productivity; although a reduction in fishing pressure, especially from bottom trawlers, is essential for the long-term sustainable use of these sharks.

Emergent community architecture despite distinct diversity in the global whale shark (Rhincodon typus) epidermal microbiome.

Microbiomes confer beneficial physiological traits to their host, but microbial diversity is inherently variable, challenging the relationship between microbes and their contribution to host health. Here, we compare the diversity and architectural complexity of the epidermal microbiome from 74 individual whale sharks (Rhincodon typus) across five aggregations globally to determine if network properties may be more indicative of the microbiome-host relationship. On the premise that microbes are expected to exhibit biogeographic patterns globally and that distantly related microbial groups can perform similar functions, we hypothesized that microbiome co-occurrence patterns would occur independently of diversity trends and that keystone microbes would vary across locations. We found that whale shark aggregation was the most important factor in discriminating taxonomic diversity patterns. Further, microbiome network architecture was similar across all aggregations, with degree distributions matching Erdos-Renyi-type networks. The microbiome-derived networks, however, display modularity indicating a definitive microbiome structure on the epidermis of whale sharks. In addition, whale sharks hosted 35 high-quality metagenome assembled genomes (MAGs) of which 25 were present from all sample locations, termed the abundant 'core'. Two main MAG groups formed, defined here as Ecogroup 1 and 2, based on the number of genes present in metabolic pathways, suggesting there are at least two important metabolic niches within the whale shark microbiome. Therefore, while variability in microbiome diversity is high, network structure and core taxa are inherent characteristics of the epidermal microbiome in whale sharks. We suggest the host-microbiome and microbe-microbe interactions that drive the self-assembly of the microbiome help support a functionally redundant abundant core and that network characteristics should be considered when linking microbiomes with host health.

Direct measurement of cruising and burst swimming speeds of the shortfin mako shark (Isurus oxyrinchus) with estimates of field metabolic rate.

The shortfin mako shark is a large-bodied pursuit predator thought to be capable of the highest swimming speeds of any elasmobranch and potentially one of the highest energetic demands of any marine fish. Nonetheless, few direct speed measurements have been reported for this species. Here, animal-borne bio-loggers attached to two mako sharks were used to provide direct measurements of swimming speeds, kinematics and thermal physiology. Mean sustained (cruising) speed was 0.90 m s-1 (±0.07 s.d.) with a mean tail-beat frequency (TBF) of 0.51 Hz (±0.16 s.d.). The maximum burst speed recorded was 5.02 m s-1 (TBFmax = 3.65 Hz) from a 2 m long female. Burst swimming was sustained for 14 s (mean speed = 2.38 m s-1 ), leading to a 0.24°C increase in white muscle temperature in the 12.5 min after the burst. Routine field metabolic rate was estimated at 185.2 mg O2 kg-1 h-1 (at 18°C ambient temperature). Gliding behaviour (zero TBF) was more frequently observed after periods of high activity, especially after capture when internal (white muscle) temperature approached 21°C (ambient temperature: 18.3°C), indicating gliding probably functions as an energy recovery mechanism and limits further metabolic heat production. The results show shortfin mako sharks generally cruise at speeds similar to other endothermic fish - but faster than ectothermic sharks - with the maximum recorded burst speed being among the highest so far directly measured among sharks, tunas and billfishes. This newly recorded high-oxygen-demand performance of mako sharks suggests it may be particularly vulnerable to habitat loss due to climate-driven ocean deoxygenation.

Evidence of increasing juvenile white sharks' (Carcharodon carcharias) habitat use at the Northern Channel Islands.

Juvenile white sharks (Carcharodon carcharias) typically aggregate along coastal beaches; however, high levels of recruitment and shifting oceanographic conditions may be causing habitat use expansions. Telemetry data indicate increased habitat use at the Northern Channel Islands (California, USA) by juvenile white shark that may be in response to increased population density at aggregation locations, or anomalous oceanographic events that impact habitat use or expand available habitat. Findings illustrate the need for long-term movement monitoring and understanding drivers of habitat use shifts and expansion to improve ecosystem management.

Tissue metal concentrations and antioxidant enzyme activity in western north Atlantic white sharks (Carcharodon carcharias).

Anthropogenic practices have increased metal contamination in marine ecosystems. Most sharks have long lifespans, occupy an important ecological position at the top of marine food webs, and can accumulate metals. However, reference levels of metal contaminants in the tissues of sharks, particularly, apex predators such as the white shark (Carcharodon carcharias), are lacking. In this study, concentrations of copper (Cu), cadmium (Cd), nickel (Ni), lead (Pb), silver (Ag), and zinc (Zn) were measured in the muscle tissue of white (n = 42) and tiger (Galeocerdo cuvier; n = 3) sharks. Metal exposure in various species, including sharks, has been correlated with increased oxidative stress. Therefore, the main objectives of this study were to assess metal accumulation and antioxidant enzyme activity (superoxide dismutase (SOD), catalase (CAT), and glutathione peroxidase (GPx)) in the muscle tissue of the population of white sharks and tiger sharks inhabiting the Western North Atlantic. The measured parameters were qualitatively compared between species. The small sample size of tiger sharks (collected from only one site) limited statistical analyses, therefore, white sharks were the primary focus of this study. Differences in tissue metal (Cu, Cd, Ni, and Zn) concentrations and antioxidant enzyme activities were detected based on collection site, with significant positive correlations between Cd and enzymes, SOD and CAT, and Zn and enzymes, SOD and GPx in C. carcharias. Differences in Ni concentration were detected based on sex, with females having higher Ni levels. Additionally, plasma osmolality was not correlated with tissue metal concentrations; however, osmolality decreased with increasing length in C. carcharias. This study is the first to report baseline levels of Cu, Zn, Cd, Ni, Ag, and Pb in muscle of North Atlantic white sharks and provides new insights into oxidative stress responses of these sensitive species to metal contaminants.

Comparison of auditory evoked potential thresholds in three shark species.

Auditory sensitivity measurements have been published for only 12 of the more than 1150 extant species of elasmobranchs (sharks, skates and rays). Thus, there is a need to further understand sound perception in more species from different ecological niches. In this study, the auditory evoked potential (AEP) technique was used to compare hearing abilities of the bottom-dwelling New Zealand carpet shark (Cephaloscyllium isabellum) and two benthopelagic houndsharks (Triakidae), the rig (Mustelus lenticulatus) and the school shark (Galeorhinus galeus). AEPs were measured in response to tone bursts (frequencies: 80, 100, 150, 200, 300, 450, 600, 800 and 1200 Hz) from an underwater speaker positioned 55 cm in front of the shark in an experimental tank. AEP detection thresholds were derived visually and statistically, with statistical measures slightly more sensitive (∼4 dB) than visual methodology. Hearing abilities differed between species, mainly with respect to bandwidth rather than sensitivity. Hearing was least developed in the benthic C. isabellum [upper limit: 300 Hz, highest sensitivity: 100 Hz (82.3±1.5 dB re. 1 µm s-2)] and had a wider range in the benthopelagic rig and school sharks [upper limit: 800 Hz; highest sensitivity: 100 Hz (79.2±1.6 dB re. 1 µm s-2) for G. galeus and 150 Hz (74.8±1.8 dB re. 1 µm s-2) for M. lenticulatus]. The data are consistent with those known for 'hearing non-specialist' teleost fishes that detect only particle motion, not pressure. Furthermore, our results provide evidence that benthopelagic sharks exploit higher frequencies (max. 800 Hz) than some of the bottom-dwelling sharks (max. 300 Hz). Further behavioural and morphological studies are needed to identify what ecological factors drive differences in upper frequency limits of hearing in elasmobranchs.

Endothermic physiology of extinct megatooth sharks.

The evolution of the extinct megatooth shark, Otodus megalodon, and its close phylogenetic relatives remains enigmatic. A central question persists regarding the thermophysiological origins of these large predatory sharks through geologic time, including whether O. megalodon was ectothermic or endothermic (including regional endothermy), and whether its thermophysiology could help to explain the iconic shark's gigantism and eventual demise during the Pliocene. To address these uncertainties, we present unique geochemical evidence for thermoregulation in O. megalodon from both clumped isotope paleothermometry and phosphate oxygen isotopes. Our results show that O. megalodon had an overall warmer body temperature compared with its ambient environment and other coexisting shark species, providing quantitative and experimental support for recent biophysical modeling studies that suggest endothermy was one of the key drivers for gigantism in O. megalodon and other lamniform sharks. The gigantic body size with high metabolic costs of having high body temperatures may have contributed to the vulnerability of Otodus species to extinction when compared to other sympatric sharks that survived the Pliocene epoch.

Intraspecific variation in muscle growth of two distinct populations of Port Jackson sharks under projected end-of-century temperatures.

Although pervasive, the effects of climate change vary regionally, possibly resulting in differential behavioral, physiological, and/or phenotypic responses among populations within broadly distributed species. Juvenile Port Jackson sharks (Heterodontus portusjacksoni) from eastern and southern Australia were reared at their current (17.6 °C Adelaide, South Australia [SA]; 20.6 °C Jervis Bay, New South Wales [NSW]) or projected end-of-century (EOC) temperatures (20.6 °C Adelaide, SA; 23.6 °C Jervis Bay, NSW) and assessed for morphological features of skeletal muscle tissue. Nearly all skeletal muscle properties including cellularity, fiber size, myonuclear domain, and satellite cell density did not differ between locations and thermal regimes. However, capillary density was significantly influenced by thermal treatment, where Adelaide sharks raised at current temperatures had a lower capillarity than Jervis Bay sharks raised at ambient or projected EOC temperatures. This may indicate higher metabolic costs at elevated temperatures. However, our results suggest that regardless of the population, juvenile Port Jackson sharks may have limited acclimatory potential to alter muscle metabolic features under a temperature increase, which may make this species vulnerable to climate change.

Free-diving sharks.

Hammerhead sharks hold their breath when diving to regulate body temperature.

"Breath holding" as a thermoregulation strategy in the deep-diving scalloped hammerhead shark.

Fish moving between different thermal environments experience heat exchange via conduction through the body wall and convection from blood flow across the gills. We report a strategy of preventing convective heat loss at the gills during excursions into deep, cold water by the tropical scalloped hammerhead shark (Sphryna lewini). Adult scalloped hammerhead sharks dive rapidly and repeatedly from warm (~26°C) surface waters to depths exceeding 800 meters with temperatures as low as 5°C. Biologgers attached to adult sharks show that warm muscle temperatures were maintained throughout the deepest portion of each dive. Substantive cooling only occurred during the latter stages of the ascent phase and, once initiated, was rapid. Heat transfer coefficient modeling indicated that convective heat transfer was suspended, probably by suppressing gill function during deep dives. This previously unobserved strategy has broad similarities to marine mammal "breath hold" diving.

Behaviour of mesopredatory coral reef fishes in response to threats from sharks and humans.

Both sharks and humans present a potentially lethal threat to mesopredatory fishes in coral reef systems, with implications for both population dynamics and the role of mesopredatory fishes in reef ecosystems. This study quantifies the antipredator behaviours mesopredatory fishes exhibit towards the presence of large coral reef carnivores and compares these behavioural responses to those elicited by the presence of snorkelers. Here, we used snorkelers and animated life-size models of the blacktip reef shark (Carcharhinus melanopterus) to simulate potential predatory threats to mesopredatory reef fishes (lethrinids, lutjanids, haemulids and serranids). The responses of these reef fishes to the models and the snorkelers were compared to those generated by three non-threatening controls (life-size models of a green turtle [Chelonia mydas], a PVC-pipe [an object control] and a Perspex shape [a second object control]). A Remote Underwater Stereo-Video System (Stereo-RUV) recorded the approach of the different treatments and controls and allowed accurate measurement of Flight Initiation Distance (FID) and categorization of the type of flight response by fishes. We found that mesopredatory reef fishes had greater FIDs in response to the approach of threatening models (1402 ± 402-1533 ± 171 mm; mean ± SE) compared to the controls (706 ± 151-896 ± 8963 mm). There was no significant difference in FID of mesopredatory fishes between the shark model and the snorkeler, suggesting that these treatments provoked similar levels of predator avoidance behaviour. This has implications for researchers monitoring behaviour in situ or using underwater census as a technique to estimate the abundance of reef fishes. Our study suggests that, irrespective of the degree to which sharks actually consume these mesopredatory reef fishes, they still elicit a predictable and consistent antipredator response that has the potential to create risk effects.

Operant conditioning as a tool to assess hearing abilities in sharks.

Sharks (elasmobranchs) are an ancient, diverse group of fishes, representing a basal stage in the evolution of vertebrate hearing. Yet, our understanding of behavioural measures of hearing abilities in sharks is limited. To address this, an operant conditioning paradigm was designed, and scalloped hammerhead Sphyrna lewini and rig (spotted estuary smooth hound) Mustelus lenticulatus were successfully trained to respond to pure-tone acoustic stimuli from an underwater speaker. After 2-3 weeks of training, both species showed distinctive responses to these acoustic stimuli and retained this behaviour when reinforced. S. lewini responded to a 400 Hz pulsed tone with an abrupt increase in tailbeat frequency (97 beats per 30 s vs. 69 beats for a 2 kHz control and 70 beats for no signal) and sustained vigorous swimming (arousal response) for at least 30 s. In response to a 200 Hz pulsed tone, M. lenticulatus visited a target area under the speaker significantly more frequently (13.4 ± 4.3 times per minute vs. 1.4 ± 1.5 times for a 1.2 kHz control and 0.9 ± 0.01 times for no signal) and swam circles under the speaker to search for food. The authors used S. lewini arousal responses to pure-tone stimuli of 40, 80, 200, 400, 600 and 800 Hz to generate a provisional hearing-threshold curve. The results show that S. lewini adapts to low-frequency hearing (greatest sensitivity at 200 Hz, upper limit 800 Hz), which is like other coastal pelagic sharks that have been investigated so far. Despite challenges operant acoustic conditioning studies are a viable method for revealing auditory capabilities of sharks.

First record of the cirripede Anelasma squalicola Darwin, 1852 (Pollicipedidae) in the Mediterranean Sea.

Anelasma squalicola Darwin 1852, is a barnacle that occurs on elasmobranchs from the Atlantic, Pacific, Arctic and Indian Ocean. It infects sharks of the families Etmopteridae and Pentanchidae. The barnacle attaches sub-dermally to its hosts, relying on its interstitial fluid for its nourishment. Herein we present the first record of this species in the Mediterranean Sea, from Etmopterus spinax (Linnaeus, 1758) caught during a bottom-trawl survey in the Alboran Sea (western Mediterranean) in 2017. The overall prevalence of infection was very low (0.22%). The calculated HSI index for the parasitized individual was lower compared to overall means of other individuals and could indicate a potential physiological alteration in the host.