Enameloid-bound δ15 N reveals large trophic separation among Late Cretaceous sharks in the northern Gulf of Mexico.

The nitrogen isotopic composition (15 N/14 N ratio, or δ15 N) of enameloid-bound organic matter (δ15 NEB ) in shark teeth was recently developed to investigate the biogeochemistry and trophic structures (i.e., food webs) of the ancient ocean. Using δ15 NEB , we present the first nitrogen isotopic evidence for trophic differences between shark taxa from a single fossil locality. We analyze the teeth of four taxa (Meristodonoides, Ptychodus, Scapanorhynchus, and Squalicorax) from the Late Cretaceous (83-84 Ma) Trussells Creek site in Alabama, USA, and compare the N isotopic findings with predictions from tooth morphology, the traditional method for inferring shark paleo-diets. Our δ15 NEB data indicate two distinct trophic groups, with averages separated by 6.1 ± 2.1‰. The lower group consists of Meristodonoides and Ptychodus, and the higher group consists of Scapanorhynchus and Squalicorax (i.e., lamniforms). This δ15 NEB difference indicates a 1.5 ± 0.5 trophic-level separation between the two groups, a finding that is in line with paleontological predictions of a higher trophic level for these lamniforms over Meristodonoides and Ptychodus. However, the δ15 NEB of Meristodonoides is lower than suggested by tooth morphology, although consistent with mechanical tests suggesting that higher trophic-level bony fishes were not a major component of their diet. Further, δ15 NEB indicates that the two sampled lamniform taxa fed at similar trophic levels despite their different inferred tooth functions. These two findings suggest that tooth morphology alone may not always be a sufficient indicator of dietary niche. The large trophic separation revealed by the δ15 NEB offset leaves open the possibility that higher trophic-level lamniforms, such as those measured here, preyed upon smaller, lower trophic-level sharks like Meristodonoides.

Relationships in Shark Skin: Mechanical and Morphological Properties Vary between Sexes and among Species.

Shark skin is a composite of mineralized dermal denticles embedded in an internal collagen fiber network and is sexually dimorphic. Female shark skin is thicker, has greater denticle density and denticle overlap compared to male shark skin, and denticle morphology differs between sexes. The skin behaves with mechanical anisotropy, extending farther when tested along the longitudinal (anteroposterior) axis but increasing in stiffness along the hoop (dorsoventral or circumferential) axis. As a result, shark skin has been hypothesized to function as an exotendon. This study aims to quantify sex differences in the mechanical properties and morphology of shark skin. We tested skin from two immature male and two immature female sharks from three species (bonnethead shark, Sphyrna tiburo; bull shark, Carcharhinus leucas; silky shark, Carcharhinus falciformis) along two orientations (longitudinal and hoop) in uniaxial tension with an Instron E1000 at a 2 mm s-1 strain rate. We found that male shark skin was significantly tougher than female skin, although females had significantly greater skin thickness compared to males. We found skin in the hoop direction was significantly stiffer than the longitudinal direction across sexes and species, while skin in the longitudinal direction was significantly more extensible than in the hoop direction. We found that shark skin mechanical behavior was impacted by sex, species, and direction, and related to morphological features of the skin.

Ontogenetic morphometry of the brown smoothhound shark Mustelus henlei with implications for ecology and evolution.

The central tenet of ecomorphology links ecological and morphological variation through the process of selection. Traditionally used to rationalise morphological differences between taxa, an ecomorphological approach is increasingly being utilised to study morphological differences expressed through ontogeny. Elasmobranchii (sharks, rays and skates) is one clade in which such ontogenetic shifts in body form have been reported. Such studies are limited to a relatively small proportion of total elasmobranch ecological and morphological diversity, and questions remain regarding the extent to which ecological selection are driving observed morphometric trends. In this study, we report ontogenetic growth trajectories obtained via traditional linear morphometrics from a large data set of the brown smoothhound shark (Mustelus henlei). We consider various morphological structures including the caudal, dorsal and pectoral fins, as well as several girth measurements. We use an ecomorphological approach to infer the broad ecological characteristics of this population and refine understanding of the selective forces underlying the evolution of specific morphological structures. We suggest that observed scaling trends in M. henlei are inconsistent with migratory behaviour, but do not contradict a putative trophic niche shift. We also highlight the role of predation pressure and sex-based ecological differences in driving observed trends in morphometry, a factor which has previously been neglected when considering the evolution of body form in sharks.

Leucism in sharks: a histological examination.

Isolated cases of skin pigment disorders, including leucism, in sharks and rays have been reported for multiple species. Nonetheless, the morphological basis behind these chromatic anomalies has not been examined histologically. In this study, the authors quantified and compared the presence of melanin in multiple tissue samples of leucistic and fully pigmented blacktip sharks Carcharhinus limbatus. The authors' results support lack of melanin to be responsible for leucistic colouration. The histological differences responsible were evaluated.

What came first, the shark or the egg? Discovery of a new species of deepwater shark by investigation of egg case morphology.

Apristurus ovicorrugatus, a new species of deepwater catshark, is described from northwestern Australia. Unique egg cases belonging to an unknown species of Apristurus prompted a more detailed investigation of Apristurus specimens off northwestern Australia. One specimen previously identified as A. sinensis collected off Dampier Archipelago was found gravid with a single egg case. Removal of this egg case confirmed that this species was responsible for producing the unique egg cases previously recorded. The egg cases of this species have strong T-shaped longitudinal ridges on the dorsal and ventral surfaces which are unique in the genus Apristurus. The ridges most closely resemble those present in Bythaelurus canescens from South America, but are larger and always T-shaped. The holotype is closest morphologically to A. sinensis but differs in having a medium brown buccal cavity (vs. jet black), ridged egg cases (vs. smooth egg cases), fewer intestinal spiral valve turns and larger pectoral fins. The holotype is also similar, and closest on a molecular level, to A. nakayai with which it shares a unique synapomorphic character, the white shiny iris (apomorphic within the genus). A late-term embryo removed from an egg case superficially resembled the holotype except in having two parallel rows of enlarged dermal denticles on the dorsolateral predorsal surface. Recent nomenclatural changes to the genera Apristurus and Pentanchus are discussed and challenged. This study highlights the important contribution that egg case morphology has on oviparous elasmobranch taxonomy.

The importance of the appendicular skeleton for the phylogenetic reconstruction of lamniform sharks (Chondrichthyes: Elasmobranchii).

Lamniform sharks are one of the more conspicuous groups of elasmobranchs, including several emblematic taxa as the white shark. Although their monophyly is well supported, the interrelationships of taxa within Lamniformes remains controversial because of the conflict among various previous molecular-based and morphology-based phylogenetic hypotheses. In this study, we use 31 characters related to the appendicular skeleton of lamniforms and demonstrate their ability to resolve the systematic interrelationships within this shark order. In particular, the new additional skeletal characters resolve all polytomies that were present in previous morphology-based phylogenetic analyses of lamniforms. Our study demonstrates the strength of incorporating new morphological data for phylogenetic reconstructions.

Contrast-enhanced computed tomography and cross-sectional anatomy of the trunk in the brownbanded bamboo shark (Chiloscyllium punctatum).

Computed tomography (CT) is used in veterinary medicine for the diagnosis of bones and soft tissue diseases in various species. In addition, CT has recently been used to diagnose aquatic animals, including Selachimorpha, which are difficult to diagnose out of water. However, because Selachimorpha do not have adipose tissue in the coelomic cavity, the coelomic organs cannot be fully identified using non-contrast CT (NCCT). The aim of this study is to present the anatomical features of the cadaver, NCCT, and contrast-enhanced CT (CECT) as well as the change in CT values of the coelomic organs and musculature of the brownbanded bamboo shark. NCCT scans were performed under anaesthesia in one male and one female shark. CECT was performed 30 min after iopamidol was administered intravenously. The sharks were euthanized, frozen at -20°C, and sliced in the same position in which they were scanned. Using electric band saw, 10-mm transversal sections were obtained. The anatomical structures of both males and females were identified by transversal sections, and CT images homologous to transversal sections were then selected. Sagittal and coronal CECT images were also obtained to facilitate understanding of the location and size of coelomic organs. Although bone structure and air in organs could be sufficiently discriminated on NCCT image, the coelomic organs were almost indistinguishable. On the other hand, CECT images obtained sufficient contrast to identify most coelomic organs in addition to bone and air. The results provide an atlas of a cross-sectional anatomy and CECT images, which is useful information for the medical diagnosis of coelomic organs in live Selachimorpha.

Head anatomy of a lantern shark wet-collection specimen (Chondrichthyes: Etmopteridae).

In this study, we apply a two-step (untreated and soft tissue stained) diffusible iodine-based contrast-enhanced micro-computed tomography array to a wet-collection Lantern Shark specimen of Etmopterus lucifer. The focus of our scanning approach is the head anatomy. The unstained CT data allow the imaging of mineralized (skeletal) tissue, while results for soft tissue were achieved after staining for 120 h in a 1% ethanolic iodine solution. Three-dimensional visualization after the segmentation of hard as well as soft tissue reveals new details of tissue organization and allows us to draw conclusions on the significance of organs in their function. Outstanding are the ampullae of Lorenzini for electroreception, which appear as the dominant sense along with the olfactory system. Corresponding brain areas of these sensory organs are significantly enlarged as well and likely reflect adaptations to the lantern sharks' deep-sea habitat. While electroreception supports the capture of living prey, the enlarged olfactory system can guide the scavenging of these opportunistic feeders. Compared to other approaches based on the manual dissection of similar species, CT scanning is superior in some but not all aspects. For example, fenestrae of the cranial nerves within the chondrocranium cannot be identified reflecting the limitations of the method, however, CT scanning is less invasive, and the staining is mostly reversible and can be rinsed out.

Jaw mechanics in macrophagous lamniform sharks and their evolutionary and functional implications.

Jaw mechanics of lamniform sharks were examined three-dimensionally to analyze the variability in jaw shape and the evolution of the jaw system based on the extant macrophagous species. Three-dimensional lever analysis was applied to lamniform jaws to calculate bite force at each tooth relative to maximum input force from jaw adductor muscles for interspecific comparison of efficiency in lamniform jaws. When total input force from the jaw adductor muscles on both working and balancing sides of the skull is considered, input force varies along the jaw because the contribution by balancing side muscles is not constant. The phylogenetically basal-most species, Mitsukurina owstoni, has the least efficient jaws due to posteriorly positioned jaw adductor muscles. Our study shows that the higher efficiency of jaws is regarded as apomorphic in lamniform phylogeny owing to the anterior extension of jaw adductor muscles relative to M. owstoni and a relative decrease in jaw length in relation to width seen in some species, both of which increase leverage. Differences in the efficiency of jaws among derived genera or species are due to the morphology of their jaws. The relationship between calculated bite force relative to maximum input force and tooth morphology indicates low relative bite forces being exerted at anteriorly located, narrow, piercing teeth, whereas high relative bite forces at posteriorly located, broad, cutting, or crushing-type teeth. As a result, the biting pressure during feeding is maintained throughout the tooth series.

Hexabothriidae and Monocotylidae (Monogenoidea) from the gills of neonate hammerhead sharks (Sphyrnidae) Sphyrna gilberti, Sphyrna lewini and their hybrids from the western North Atlantic Ocean.

Neonates of hammerhead sharks (Sphyrnidae), Sphyrna lewini (Griffith and Smith, 1834), the sympatric cryptic species, Sphyrna gilberti Quattro et al., 2013, and their hybrids were captured in the western North Atlantic, along the coast of South Carolina, USA, between 2018 and 2019 and examined for gill monogenoids. Parasites were identified and redescribed from the gills of 79 neonates, and DNA sequences from partial fragments of the nuclear 28S ribosomal RNA (rDNA) and cytochrome c oxidase I mitochondrial DNA (COI) genes were generated to confirm species identifications. Three species of monogenoids from Hexabothriidae Price, 1942 and Monocotylidae Taschenberg, 1879 were determined and redescribed. Two species of Hexabothriidae, Erpocotyle microstoma (Brooks, 1934) and Erpocotyle sphyrnae (MacCallum, 1931), infecting both species of Sphyrna and hybrids; and 1 species of Monocotylidae, Loimosina wilsoni Manter, 1944, infecting only S. lewini and hybrids. Loimosina wilsoni 28S rDNA sequences matched those of Loimosina sp. from the southern coast of Brazil. Based on limited morphological analysis, Loimosina parawilsoni is likely a junior synonym of L. wilsoni. This is the first taxonomic study of monogenoids infecting S. gilberti and hybrids of S. gilberti and S. lewini.

Biomechanics of the jaws of spotted ratfish.

Elasmobranch fishes (sharks, skates and rays) consume prey of a variety of sizes and properties, and the feeding mechanism typically reflects diet. Spotted ratfish, Hydrolagus colliei (Holocephali, sister group of elasmobranchs), consume both hard and soft prey; however, the morphology of the jaws does not reflect the characteristics typical of durophagous elasmobranchs. This study investigated the mechanical properties and morphological characteristics of the jaws of spotted ratfish over ontogeny, including strain, stiffness and second moment of area, to evaluate the biomechanical function of the feeding structures. Compressive stiffness of the jaws (E=13.51-21.48 MPa) is similar to that of silicone rubber, a very flexible material. In Holocephali, the upper jaw is fused to the cranium; we show that this fusion reduces deformation experienced by the upper jaw during feeding. The lower jaw resists bending primarily in the posterior half of the jaw, which occludes with the region of the upper jaw that is wider and flatter, thus potentially providing an ideal location for the lower jaw to crush or crack prey. The mechanical properties and morphology of the feeding apparatus of spotted ratfish suggest that while the low compressive stiffness is a material limit of the jaw cartilage, spotted ratfish, and perhaps all holocephalans, evolved structural solutions (i.e. fused upper jaw, shape variation along lower jaw) to meet the demands of a durophagous diet.

Ontogenetic changes in the tooth morphology of bull sharks (Carcharhinus leucas).

Teeth are an integral component of feeding ecology, with a clear link between tooth morphology and diet, as without suitable dentition prey cannot be captured nor broken down for consumption. Bull sharks, Carcharhinus leucas, undergo an ontogenetic niche shift from freshwater to marine habitats, which raises the question: does tooth morphology change with ontogeny? Tooth shape, surface area and thickness were measured using both morphometrics and elliptic Fourier analysis to determine if morphology varied with position in the jaw and if there was an ontogenetic change concordant with this niche shift. Significant ontogenetic differences in tooth morphology as a function of position in the jaw and shark total length were found, with upper and lower jaws of bull sharks presenting two different tooth morphologies. Tooth shape and thickness fell into two groupings, anterior and posterior, in both the upper and lower jaws. Tooth surface area, however, indicated three groupings, mesial, intermediate and distal, in both the upper and lower jaws. While tooth morphology changed significantly with size, showing an inflection at sharks of 135 cm total length, each morphological aspect retained the same tooth groupings throughout. These ontogenetic differences in tooth morphologies reflect tooth strength, prey handling and heterodonty.

Morphological pattern of the dermal denticles of the Southern sawtail catshark Galeus mincaronei Soto, 2001.

Dermal denticles cover the skin of sharks for protection and may have additional functions, depending on the environment. The aim of the present study was to analyze the morphological pattern of dermal denticles across the body of the poorly known deep-water catshark Galeus mincaronei, which is endemic to Southern Brazil. Additionally, we aimed to identify differences between sexes and stages of sexual maturity. For this purpose, we analyzed patches of eight skin samples from body regions, that is, the head, body, and tail. Dermal denticles were imaged using scanning electron microscopy. Measurements of crown morphological variations (length and width) and density of dermal denticles were performed. Nine characters of dermal denticle morphology were selected to describe a morphological pattern using principal component analysis (PCA). The first PCA axis explained 45.2% of the variation, the second explained 33%, and the third explained 14.3%, together explaining 92.5% of the total variation in denticle morphology. We identified three groups of dermal denticle crown morphologies and their possible functions. The first was composed of nostril and snout denticles, which might suggest abrasion strength functions; the second was composed of most body denticles, such as those in the cranium, on the left side of the body, and on the pectoral, dorsal, and caudal fins, which might be related to generalized functions; and the third was composed of only caudal crest denticles, which might indicate defense functions with hydrodynamic properties in G. mincaronei. Likewise, our results reveal variations in dermal denticle morphology and arrangement along the three maturity stages considered in both sexes. Similarly, sexual dimorphism in the crown width and morphology of dermal denticles was observed in developing and mature individuals. Our results are the first to reveal the morphological variation of dermal denticles in relation to function, sex, and ontogeny of individual G. mincaronei.

Sixth sense in the deep-sea: the electrosensory system in ghost shark Chimaera monstrosa.

Animals that continually live in deep sea habitats face unique challenges and require adaptive specializations solutions in order to locate and identify food, predators, and conspecifics. The Ampullae of Lorenzini are specialized electroreceptors used by chondrichthyans for important biological functions. Ampullary organs of the ghost shark Chimaera monstrosa, a deep-sea species commonly captured as by-catch in the bottom trawl fishery, are here described for the first time using macroscopic, ultrastructural and histological approaches. The number of ampullary pores in C. monstrosa is about 700, distributed into the whole cephalic section of C. monstrosa, and organized in12 pore clusters and they are arranged into different configurations and form a distinct morphological pattern for this species, showing some anatomical peculiarities never described before in others cartilaginous fishes and may constitute an evolutionary adaptation of this ancient chondrichthyan species to the extreme environmental conditions of its deep sea niche.

Morphological study of the oral denticles of the porbeagle shark Lamna nasus.

Oral denticles of sharks are composed by a crown, dentine covered by a layer of enameloid and pulp cavity, the same structure of the dermal denticles found across the body surface of most elasmobranchs. In addition, oral papillae and taste buds are distributed among denticles within the oropharyngeal cavity, playing a fundamental role for tasting as part of the chemosensory system of fishes. Scanning electron microscopy (SEM) has been employed as an important tool for the study of dermal denticles and other structures, as well as histology and more recently computed tomography (CT) scan analysis. Herein, the authors used two methods for the study of the morphology of the oropharyngeal cavity of Lamna nasus (Lamniformes), an oceanic and pelagic shark: SEM and CT scan. The general morphology of oral denticles studied herein is related to abrasion strength as they are diamond-shaped, lack lateral cusps and have less pronounced ridges. In addition, smooth ridges and broad rounded denticles could be related to prevent abrasion during food consumption and manipulation. Oral papillae had a round shape and were observed only under SEM. The densities of papillae were estimated in 100 per cm2 , whereas denticles were 1760 and 1230 cm2 over the dorsal and ventral regions, respectively. The high numbers of denticles are inversely proportional to papillae density; denticles seem to restrict papillae distribution. Regarding the differences between methodologies, under SEM, only the crown was visualized, as well the papillae, allowing the estimation of size and density of both structures. Nonetheless, under CT scan, the whole components of denticles were clearly visualized: different views of the crown, peduncle, basal plate, and pulp cavity. On the contrary, oral papillae were not visualized under CT due to the tissue preparation. Furthermore, both methods are complementary and were important to extract as much information as possible from denticles and papillae.

Dental pathologies in lamniform and carcharhiniform sharks with comments on the classification and homology of double tooth pathologies in vertebrates.

Double tooth pathologies are important indicators of trauma, disease, diet, and feeding biomechanics, and are widely documented in mammals. However, diagnosis of double tooth pathologies in extinct non-mammalian vertebrates is complicated by several compounding factors including: a lack of shared terminology reflecting shared etiology, inconsistencies in definitions and key features within and outside of mammals (e.g., gemination, fusion, twinning, concrescence); differences in tooth morphology, heterodonty, regeneration, and implantation between mammals and non-mammalian vertebrates; and the unmet need for diagnostic criteria that can be applied to isolated teeth, which are common in the fossil record. Here we report on double tooth pathologies in the lamniform and carcharhiniform Cenozoic sharks Otodus megalodon (NCSM 33639) and Carcharhinus leucas (NCSM 33640, 33641). All three teeth bear a singular bifid crown with mirrored halves and abnormal internal microstructure-a single, bifurcating pulp cavity in C. leucas and a more than tripling of vessels in O. megalodon (from two to seven main ascending canals). We identify these abnormalities as likely examples of gemination due to their symmetry, which rules out fusion of tooth buds in one tooth file in different developmental stages in polyphyodont taxa; however, we note that incomplete forms of mesiodistal tooth fusion can be morphologically indistinguishable from gemination, and thus fusion cannot be rejected. We further compile and recategorize, when possible, the diversity of tooth pathologies in sharks. The identification of double tooth pathologies in O. megalodon and C. leucas has paleobiological implications. Such pathologies in sharks are largely hypothesized to stem from trauma to developing tooth buds. Carcharhinus leucas is known to feed on prey documented to cause feeding-related oral traumas (e.g., rays, sawfish, spiny fish, and sea urchins). However, O. megalodon, is considered to have largely fed on marine mammals, and perhaps turtles and/or fish, raising the possibility that the dietary diversity of this species is, as of yet, underappreciated. The genetic underpinnings of tooth morphogenesis and regeneration is highly conserved throughout vertebrate evolution, suggesting a homologous framework can be established. However, more research is needed to link developmental, paleobiological, and/or paleoenvironmental factors to gemination/fusion in polyphyodont taxa. We argue that the definitions and diagnostic criteria for dental pathologies in vertebrates require standardization in order to advance macroevolutionary studies of feeding trauma in deep time.

Heterodonty and ontogenetic shift dynamics in the dentition of the tiger shark Galeocerdo cuvier (Chondrichthyes, Galeocerdidae).

The lifelong tooth replacement in elasmobranch fishes (sharks, rays and skates) has led to the assemblage of a great number of teeth from fossil and extant species, rendering tooth morphology an important character for taxonomic descriptions, analysing phylogenetic interrelationships and deciphering their evolutionary history (e.g. origination, divergence, extinction). Heterodonty (exhibition of different tooth morphologies) occurs in most elasmobranch species and has proven to be one of the main challenges for these analyses. Although numerous shark species are discovered and described every year, detailed descriptions of tooth morphologies and heterodonty patterns are lacking or are only insufficiently known for most species. Here, we use landmark-based 2D geometric morphometrics on teeth of the tiger shark Galeocerdo cuvier to analyse and describe dental heterodonties among four different ontogenetic stages ranging from embryo to adult. Our results reveal rather gradual and subtle ontogenetic shape changes, mostly characterized by increasing size and complexity of the teeth. We furthermore provide the first comprehensive description of embryonic dental morphologies in tiger sharks. Also, tooth shapes of tiger sharks in different ontogenetic stages are re-assessed and depicted in detail. Finally, multiple cases of tooth file reversal are described. This study, therefore, contributes to our knowledge of dental traits across ontogeny in the extant tiger shark G. cuvier and provides a baseline for further morphological and genetic studies on the dental variation in sharks. Therefore, it has the potential to assist elucidating the underlying developmental and evolutionary processes behind the vast dental diversity observed in elasmobranch fishes today and in deep time.

Shark centra microanatomy and mineral density variation studied with laboratory microComputed Tomography.

Centra of shark vertebrae from three species of Lamniformes (Alopias vulpinus, Carcharodon carcharias and Isurus oxyrinchus) and three species of Carcharhiniformes (Carcharhinus plumbeus, Carcharhinus obscurus and Prionace glauca) were imaged with laboratory microcomputed Tomography (microCT) using volume element (voxel) sizes between 16 and 24 µm. Linear attenuation coefficients were the same in the corpus calcarea (hour-glass-shaped cone) and intermedialia of the lamniforms but were smaller in the intermedialia than in the corpus calcarea of the carcharhiniforms. All centra contained growth bands which were visible as small changes in linear attenuation coefficient. In all six cases, the cross-sections of the cones were close to circular, and the cone angles matched those reported in the literature. Cartilage canals were a prominent structure in the intermedialia of all species, 3D renderings of centra of C. obscurus and I. oxyrinchus diameters showed these canals ran radially outward from the cone walls, and canal diameters were consistent with the limited numerical values in the literature. Somewhat higher calcification levels around the periphery of cartilage canals and of outer surfaces of the intermedialia and corpus calcerea suggest microstructural variation exists at scale below that which can be resolved in the present data sets.

Branching patterns of the afferent branchial arteries and their phylogenetic significance in rays (Batoidea).

Rays of the superorder Batoidea comprise the most diverse group of chondrichthyans in terms of valid species and morphological disparity. Up to the present little agreement is observed in studies based on morphological and molecular data focused on uncovering the interrelationships within Batoidea. Morphology-based phylogenies of batoids have not included characters related to the afferent branchial arteries, and little is known about the variation in this anatomical complex in rays. Herein, representatives of 32 genera from 19 families currently recognized of rays were examined as well as some shark taxa. Seven new characters are proposed and tested in two different analyses, one on their own and in the other they were added to the morphological data matrix of the most recent analysis of interrelationships within Batoidea. The arrangement of afferent branchial arteries differs mainly among orders and families of batoids. The absence of a common trunk from which the three posteriormost afferent arteries branch is interpreted as a synapomorphy for Myliobatiformes and the presence of a coronary cranial artery as an autapomorphy for Mobula hypostoma. A close spatial relationship between the second and third afferent arteries within the common branch from the ventral aorta is proposed as a synapomorphy for Rajiformes with a secondary modification in Sympterygia. Data about patterns in afferent branchial arteries in additional taxa such as Squaliformes and Chimaeriformes are needed to better understand the evolution of this character complex among chondrichthyans.

Feeding ecology has shaped the evolution of modern sharks.

Sharks are iconic predators in today's oceans, yet their modern diversity has ancient origins. In particular, present hypotheses suggest that a combination of mass extinction, global climate change, and competition has regulated the community structure of dominant mackerel (Lamniformes) and ground (Carcharhiniformes) sharks over the last 66 million years. However, while these scenarios advocate an interplay of major abiotic and biotic events, the precise drivers remain obscure. Here, we focus on the role of feeding ecology using a geometric morphometric analysis of 3,837 fossil and extant shark teeth. Our results reveal that morphological segregation rather than competition has characterized lamniform and carcharhiniform evolution. Moreover, although lamniforms suffered a long-term disparity decline potentially linked to dietary "specialization," their recent disparity rivals that of "generalist" carcharhiniforms. We further confirm that low eustatic sea levels impacted lamniform disparity across the end-Cretaceous mass extinction. Adaptations to changing prey availability and the proliferation of coral reef habitats during the Paleogene also likely facilitated carcharhiniform dispersals and cladogenesis, underpinning their current taxonomic dominance. Ultimately, we posit that trophic partitioning and resource utilization shaped past shark ecology and represent critical determinants for their future species survivorship.