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Shark cartilage presents a complex material composed of collagen, proteoglycans, and bioapatite. In the present study, we explored the link between microstructure, chemical composition, and biomechanical function of shark vertebral cartilage using Polarized Light Microscopy (PLM), Atomic Force Microscopy (AFM), Confocal Raman Microspectroscopy, and Nanoindentation. Our investigation focused on vertebrae from Blacktip and Shortfin Mako sharks. As typical representatives of the orders Carcharhiniformes and Lamniformes, these species differ in preferred habitat, ecological role, and swimming style. We observed structural variations in mineral organization and collagen fiber arrangement using PLM and AFM. In both sharks, the highly calcified corpus calcarea shows a ridged morphology, while a chain-like network is present in the less mineralized intermedialia. Raman spectromicroscopy demonstrates a relative increase of glucosaminocycans (GAGs) with respect to collagen and a decrease in mineral-rich zones, underlining the role of GAGs in modulating bioapatite mineralization. Region-specific testing confirmed that intravertebral variations in mineral content and arrangement result in distinct nanomechanical properties. Local Young's moduli from mineralized regions exceeded bulk values by a factor of 10. Overall, this work provides profound insights into a flexible yet strong biocomposite, which is crucial for the extraordinary speed of cartilaginous fish in the worlds’ oceans. 

Synopsis To smell, fish rely on passive water flow into their olfactory chambers and through their olfactory rosettes to detect chemical signals in their aquatic environment. The olfactory rosette is made up of secondarily folded tissues called olfactory lamellae. The olfactory morphology of cartilaginous fishes varies widely in both rosette gross morphology and lamellar microstructure. Previous research has shown differences in lamellar sensory morphology depending on the position along the rosette in hammerheads (family Sphyrnidae). Here, we investigate if this pattern continues in members of two other chondrichthyan families: Squalidae and Chimaeridae. Using contrast-enhanced microCT and scanning electron microscopy, we investigated patterns in lamellar morphology based on lamellar position along the olfactory rosette in Pacific spiny dogfish (Squalus suckleyi) and spotted ratfish (Hydrolagus colliei). We describe the gross olfactory rosette anatomy and lamellar microstructure of both species. We also put forth a new method, combining 3D morphological microCT data with 2D SEM microstructure data to better approximate lamellar sensory surface area. We found that in both species, lamellae in the center of the rosette were larger with more secondary folds. However, we found no significant differences in lamellar sensory surface area among lamellar positions. Previously, differences in lamellar sensory morphology have been tied to the internal fluid dynamics of the olfactory chamber. It is possible that the internal flow dynamics of these species are like other chondrichthyan models, where water flow patterns differ in the lateral vs the medial part of the organ, and the consistent distribution of sensory tissue does not correspond to this flow. Alternatively, the olfactory morphology of these species may result in uniform flow patterns throughout the olfactory chamber, correlating with the consistent distribution of sensory tissue throughout the organ. This study emphasizes that further investigations into chondrichthyan fluid dynamics is paramount to any future studies on the correlations between distribution of sensory tissues and water flow. 

Thresher sharks (Alopiasspp.) are characterized by an elongated, scythe-like caudal fin that is used in tail-whipping, a behaviour where the tail is thrown overhead to stun prey. Tail-whipping is performed via extreme dorsoventral bending of the vertebral column, and is dramatically different from lateral oscillatory motion used for swimming. Previous work has examined thresher shark vertebral morphology and mechanical properties, but in the context of swimming loads. Our goal was to assess centra morphometrics and microarchitecture for variations that may support extreme dorsoventral bending. We examined anterior and posterior body vertebrae from an embryo, five juvenile, and four adult thresher sharks using micro-computed tomography. We used principal component and landmark analyses to examine variables influencing vertebral morphology and mineral arrangement, respectively. We found that morphology and microstructure significantly varied across body regions and ontogeny. We hypothesize that anterior body vertebrae increase stability, while posterior body vertebrae support the caudal fin. Vertebral size and quantity of mineral structures (lamellae and nodes) increased across ontogeny, suggesting vertebrae adapt over development to support a larger body and tail. Based on our results, we hypothesize that thresher shark vertebrae vary in morphometrics and mineralization (amount and arrangement) supporting the mechanical needs for tail-whipping. 

Abstract The epaulette shark, Hemiscyllium ocellatum, is a small, reef-dwelling, benthic shark that—using its paired fins—can walk, both in and out of water. Within the reef flats, this species experiences short periods of elevated CO2 and hypoxia as well as fluctuating temperatures as reef flats become isolated with the outgoing tide. Past studies have shown that this species is robust (i.e., respiratory and metabolic performance, behavior) to climate change-relevant elevated CO2 levels as well as hypoxia and anoxia tolerant. However, epaulette shark embryos reared under ocean warming conditions hatch earlier and smaller, with altered patterns and coloration, and with higher metabolic costs than their current-day counterparts. Findings to date suggest that this species has adaptations to tolerate some, but perhaps not all, of the challenging conditions predicted for the 21st century. As such, the epaulette shark is emerging as a model system to understand vertebrate physiology in changing oceans. Yet, few studies have investigated the kinematics of walking and swimming, which may be vital to their biological fitness, considering their habitat and propensity for challenging environmental conditions. Given that neonates retain embryonic nutrition via an internalized yolk sac, resulting in a bulbous abdomen, while juveniles actively forage for worms, crustaceans, and small fishes, we hypothesized that difference in body shape over early ontogeny would affect locomotor performance. To test this, we examined neonate and juvenile locomotor kinematics during the three aquatic gaits they utilize—slow-to-medium walking, fast walking, and swimming—using 13 anatomical landmarks along the fins, girdles, and body midline. We found that differences in body shape did not alter kinematics between neonates and juveniles. Overall velocity, fin rotation, axial bending, and tail beat frequency and amplitude were consistent between early life stages. Data suggest that the locomotor kinematics are maintained between neonate and juvenile epaulette sharks, even as their feeding strategy changes. Studying epaulette shark locomotion allows us to understand this—and perhaps related—species’ ability to move within and away from challenging conditions in their habitats. Such locomotor traits may not only be key to survival, in general, as a small, benthic mesopredator (i.e., movements required to maneuver into small reef crevices to avoid aerial and aquatic predators), but also be related to their sustained physiological performance under challenging environmental conditions, including those associated with climate change—a topic worthy of future investigation. 

Abstract The elongate body plan is present in many groups of fishes, and this morphology dictates functional consequences seen in swimming behavior. Previous work has shown that increasing the number of vertebrae, or decreasing the intervertebral joint length, in a fixed length artificial system increases stiffness. Tails with increased stiffness can generate more power from tail beats, resulting in an increased mean swimming speed. This demonstrates the impacts of morphology on both material properties and kinematics, establishing mechanisms for form contributing to function. Here, we wanted to investigate relationships between form and ecological function, such as differences in dietary strategies and habitat preferences among fish species. This study aims to characterize and compare the kinematics, material properties, and vertebral morphology of four species of elongate fishes: Anoplarchus insignis, Anoplarchus purpurescens, Xiphister atropurpureus, and Xiphister mucosus. We hypothesized that these properties would differ among the four species due to their differential ecological niches. To calculate kinematic variables, we filmed these fishes swimming volitionally. We also measured body stiffness by bending the abdominal and tail regions of sacrificed individuals in different stages of dissection (whole body, removed skin, removed muscle). Finally, we counted the number of vertebrae from CT scans of each species to quantify vertebral morphology. Principal component and linear discriminant analyses suggested that the elongate fish species can be distinguished from one another by their material properties, morphology, and swimming kinematics. With this information combined, we can draw connections between the physical properties of the fishes and their ecological niches. 

Recent work showed that two species of hammerhead sharks operated as a double oscillating system, where frequency and amplitude differed in the anterior and posterior parts of the body. We hypothesized that a double oscillating system would be present in a large, volitionally swimming, conventionally shaped carcharhinid shark. Swimming kinematics analyses provide quantification to mechanistically examine swimming within and among species. Here, we quantify blacktip shark (Carcharhinus limbatus) volitional swimming kinematics under natural conditions to assess variation between anterior and posterior body regions and demonstrate the presence of a double oscillating system. We captured footage of 80 individual blacktips swimming in the wild using a DJI Phantom 4 Pro aerial drone. The widespread accessibility of aerial drone technology has allowed for greater observation of wild marine megafauna. We used Loggerpro motion tracking software to track five anatomical landmarks frame by frame to calculate tailbeat frequency, tailbeat amplitude, speed, and anterior/posterior variables: amplitude and frequency of the head and tail, and the body curvature measured as anterior and posterior flexion. We found significant increases in tailbeat frequency and amplitude with increasing swimming speed. Tailbeat frequency decreased and tailbeat amplitude increased as posterior flexion amplitude increased. We found significant differences between anterior and posterior amplitudes and frequencies, suggesting a double oscillating modality of wave propagation. These data support previous work that hypothesized the importance of a double oscillating system for increased sensory perception. These methods demonstrate the utility of quantifying swimming kinematics of wild animals through direct observation, with the potential to apply a biomechanical perspective to movement ecology paradigms.