Three-dimensional shape of natural riblets in the white shark: relationship between the denticle morphology and swimming speed of sharks.

The ridges of the dermal denticles of migratory sharks have inspired riblets to reduce the frictional drag of a fluid. In particular, the dermal denticles of white sharks (Carcharodon carcharias) are characterized by a high middle ridge and low side ridges. The detailed morphology of their denticles and their variation along the body, however, have never been investigated. Moreover, the hydrodynamic function of high-low combinations of ridges is unknown. In this article, the ridge spacings and heights of the white shark denticles were three-dimensionally quantified using microfocus X-ray computed tomography. Then, the swimming speed at which the ridges would reduce drag was hydrodynamically calculated with a flat plate body model and previous riblet data. High ridges with a large spacing were found to effectively reduce drag at a migration speed of 2.3 m s-1, while adjacent high and low ridges with a small spacing reduced drag at a burst hunting speed of 5.1 m s-1. Moreover, the above hydrodynamic calculation method was also applied to the shortfin mako shark and an extinct giant shark (called megalodon) with known ridge spacings, resulting in the estimated hunting speeds of 10.5 m s-1and 5.9 m s-1, respectively.

The construction of ofloxacin detection in fish matrix based on a shark-derived single-domain antibody.

BACKGROUND: Due to the serious issue of ofloxacin (OFL) abuse, there is an increasingly urgent need for accurate and rapid detection of OFL. Immunoassay has become the "golden method" for detecting OFL in complex matrix beneficial to its applicability for a large-scale screening, rapidity, and simplicity. However, traditional antibodies used in immunoassay present challenges such as time-consuming preparation, unstable sensitivity and specificity, and difficulty in directional evolution. In this paper, we successfully developed an OFL detection method based on a shark-derived single-domain antibody (ssdAb) to address these issues. RESULTS: Using phage display technology and a heterologous expression system, OFL-specific clones 1O11, 1O13, 1O17, 1O19, 1O21, and 2O26 were successfully isolated and expressed in soluble form. Among all OFL-specific ssdAbs, the 1O17 ssdAb exhibited the highest binding affinity to OFL in a concentration-dependence manner. The limit of detection (IC10) of 1O17 ssdAb was calculated as 0.34 ng/mL with a detection range of 3.40-1315.00 ng/mL, and its cross reactivity with other analogs was calculated to be less than 5.98 %, indicating high specificity and sensitivity. Molecular docking results revealed that 100Trp and 101Arg located in the CDR3 region of 1O17 ssdAb were crucial for OFL binding. In fish matrix performance tests, the 1O17 ssdAb did not demonstrate severe matrix interference in OFL-negative fish matrix, achieving satisfactory recovery rates ranging from 83.04 % to 108.82 % with high reproducibility. SIGNIFICANCE: This research provides a new and efficient OFL detection recognition element with significant potential in immunoassay applications, broadening the application scenarios of ssdAbs. It offers valuable insights into the structure-activity relationship between ssdAbs and small molecules, laying a theoretical foundation for the further directional modification and maturation of ssdAbs in subsequent applications.

Ecological roles and importance of sharks in the Anthropocene Ocean.

In ecosystems, sharks can be predators, competitors, facilitators, nutrient transporters, and food. However, overfishing and other threats have greatly reduced shark populations, altering their roles and effects on ecosystems. We review these changes and implications for ecosystem function and management. Macropredatory sharks are often disproportionately affected by humans but can influence prey and coastal ecosystems, including facilitating carbon sequestration. Like terrestrial predators, sharks may be crucial to ecosystem functioning under climate change. However, large ecosystem effects of sharks are not ubiquitous. Increasing human uses of oceans are changing shark roles, necessitating management consideration. Rebuilding key populations and incorporating shark ecological roles, including less obvious ones, into management efforts are critical for retaining sharks' functional value. Coupled social-ecological frameworks can facilitate these efforts.

Accumulation of Per- and Polyfluoroalkyl Substances (PFAS) in Coastal Sharks from Contrasting Marine Environments: The New York Bight and The Bahamas.

Per- and polyfluoroalkyl substances (PFAS) enter the marine food web, accumulate in organisms, and potentially have adverse effects on predators and consumers of seafood. However, evaluations of PFAS in meso-to-apex predators, like sharks, are scarce. This study investigated PFAS occurrence in five shark species from two marine ecosystems with contrasting relative human population densities, the New York Bight (NYB) and the coastal waters of The Bahamas archipelago. The total detected PFAS (∑PFAS) concentrations in muscle tissue ranged from 1.10 to 58.5 ng g-1 wet weight, and perfluorocarboxylic acids (PFCAs) were dominant. Fewer PFAS were detected in Caribbean reef sharks (Carcharhinus perezi) from The Bahamas, and concentrations of those detected were, on average, ∼79% lower than in the NYB sharks. In the NYB, ∑PFAS concentrations followed: common thresher (Alopias vulpinus) > shortfin mako (Isurus oxyrinchus) > sandbar (Carcharhinus plumbeus) > smooth dogfish (Mustelus canis). PFAS precursors/intermediates, such as 2H,2H,3H,3H-perfluorodecanoic acid and perfluorooctanesulfonamide, were only detected in the NYB sharks, suggesting higher ambient concentrations and diversity of PFAS sources in this region. Ultralong-chain PFAS (C ≥ 10) were positively correlated with nitrogen isotope values (δ15N) and total mercury in some species. Our results provide some of the first baseline information on PFAS concentrations in shark species from the northwest Atlantic Ocean, and correlations between PFAS, stable isotopes, and mercury further contextualize the drivers of PFAS occurrence.

Detailed characterization of the complete mitochondrial genome of the oceanic whitetip shark Carcharhinus longimanus (Poey, 1861).

BACKGROUND: The oceanic whitetip shark Carcharhinus longimanus (family Carcharhinidae) is one of the largest sharks inhabiting all tropical and subtropical oceanic regions. Due to their life history traits and mortality attributed to pelagic longline fishing practices, this species is experiencing substantial population decline. Currently, C. longimanus is considered by the IUCN Red List of Threatened Species as "vulnerable" throughout its range and "critically endangered" in the western north Atlantic. This study sequences and describes the complete mitochondrial genome of C. longimanus in detail. METHODS AND RESULTS: The mitochondrial genome of C. longimanus was assembled through next-generation sequencing and then analyzed using specialized bioinformatics tools. The circular, double-stranded AT-rich mitogenome of C. longimanus is 16,704 bp long and contains 22 tRNA genes, 2 rRNA genes, 13 protein coding genes and a 1,065 bp long control region (CR). Out of the 22 tRNA genes, only one (tRNA-Ser1) lacked a typical 'cloverleaf' secondary structure. The prevalence of TTA (Leu), ATT (Ile) and CTA (Leu) codons in the PCGs likely contributes to the AT-rich nature of this mitogenome. In the CR, ten microsatellites were detected but no tandem repeats were found. Stem-and-loop secondary structures were common along the entire length of the CR. Ka/Ks values estimated for all PCGs were < 1, indicating that all the PCGs experience purifying selection. A phylomitogenomic analysis based on translated PCGs confirms the sister relationship between C. longimanus and C. obscurus. The analysis did not support the monophyly of the genus Carcharhinus. CONCLUSIONS: The assembled mitochondrial genome of this pelagic shark can provide insight into the phylogenetic relationships in the genus Carcharhinus and aid conservation and management efforts in the Central Pacific Ocean.

Who bit the boat? New DNA collection and genomic methods enable species identification in suspected shark-related incidents.

Species identification following shark-related incidents is critical for effective incident management and for collecting data to inform shark-bite mitigation strategies. Witness statements are not always reliable, and species identification is often ambiguous or missing. Alternative methods for species identification include morphological assessments of bite marks, analysis of collected teeth at the scene of the incident, and genetic approaches. However, access to appropriate collection media and robust genetic assays have limited the use of genetic technologies. Here, we present a case study that facilitated a unique opportunity to compare the effectiveness of medical gauze readily available in first-aid kits, and forensic-grade swabs in collecting genetic material for shark-species identification. Sterile medical gauze and forensic-grade swabs were used to collect transfer DNA from the bite margins on a bitten surf ski which were compared to a piece of shark tissue embedded along the bite margin. Witness accounts and the characteristics of the bite mark impressions inferred the involvement of a Carcharodon carcharias (white shark). The morphology of a tooth found on the boat that picked up the surf ski, however, suggested it belonged to an Orectolobus spp. (wobbegong). Genetic analysis of DNA transferred from the shark to the surf ski included the application of a broad-target nested PCR assay followed by Sanger sequencing, with white shark contribution to the 'total sample DNA' determined with a species-specific qPCR assay. The results of the genetic analyses were congruent between sampling methods with respect to species identification and the level of activity inferred by the donor-specific DNA contribution. These data also supported the inferences drawn from the bite mark morphology. DNA from the recovered tooth was PCR amplified with a wobbegong-specific primer pair designed for this study to corroborate the tooth's morphological identification. Following the confirmation of gauze used for sampling in the case study event, two additional isolated incidents occurred and were sampled in situ using gauze, as typically found in a first-aid kit, by external personnel. DNA extracted from these gauze samples resulted in the identification of a white shark as the donor of the DNA collected from the bite marks in both instances. This study, involving three incidents separated by time and location, represents the seminal application of gauze as a sampling media after critical human-shark interactions and strongly supports the practical implementation of these methods in the field.

Personal electric deterrents can reduce shark bites from the three species responsible for the most fatal interactions.

The frequency of unprovoked shark bites is increasing worldwide, leading to a growing pressure for mitigation measures to reduce shark-bite risk while maintaining conservation objectives. Personal shark deterrents are a promising and non-lethal strategy that can protect ocean users, but few have been independently and scientifically tested. In Australia, bull (Carcharhinus leucas), tiger (Galeocerdo cuvier), and white sharks (Carcharodon carcharias) are responsible for the highest number of bites and fatalities. We tested the effects of two electric deterrents (Ocean Guardian's Freedom+ Surf and Freedom7) on the behaviour of these three species. The surf product reduced the probability of bites by 54% across all three species. The diving product had a similar effect on tiger shark bites (69% reduction) but did not reduce the frequency of bites from white sharks (1% increase), likely because the electrodes were placed further away from the bait. Electric deterrents also increased the time for bites to occur, and frequency of reactions and passes for all species tested. Our findings reveal that both Freedom+ Surf and Freedom7 electric deterrents affect shark behaviour and can reduce shark-bite risk for water users, but neither product eliminated the risk of shark bites entirely. The increasing number of studies showing the ability of personal electric deterrents to reduce shark-bite risk highlights personal protection as an effective and important part of the toolbox of shark-bite mitigation measures.

"Cocaine Shark": First report on cocaine and benzoylecgonine detection in sharks.

Cocaine (COC) and benzoylecgonine (BE), the main COC metabolite, have been detected in aquatic ecosystems. Studies focusing on wild fish are, however, very limited, and no reports concerning elasmobranchs are available. This study investigated COC and BE levels in Brazilian Sharpnose sharks (Rhizoprionodon lalandii) (n = 13) using LC-MS/MS. All samples (13/13) tested positive for COC, with 92 % (12/13) testing positive for BE. COC concentrations (23.0 µg kg-1) were over 3-fold higher than BE (7.0 µg kg-1). COC levels were about three-fold significantly higher in muscle (33.8 ± 33.4 g kg-1) compared to liver (12.2 ± 14.2 µg kg-1). Females presented higher COC concentrations in muscle (40.2 ± 35.8 µg kg-1) compared to males (12.4 ± 5.9 µg kg-1). Several positive statistical correlations were noted between COC and BE (rho = 0.84) in females, indicating systemic COC transport and metabolization, as well as between BE and weight (rho = 0.62), and between COC and the Condition Factor (rho = 0.73). A strong correlation was noted between BE and COC in the muscle of non-pregnant females (rho = 1.00). This study represents the first COC and BE report in free-ranging sharks, and the findings point to the potential impacts of the presence of illicit drugs in environments.

Genome analysis reveals three distinct lineages of the cosmopolitan white shark.

The white shark (Carcharodon carcharias) (Linnaeus, 1758), an iconic apex predator occurring in all oceans,1,2 is classified as Vulnerable globally3-with global abundance having dropped to 63% of 1970s estimates,4-and as Critically Endangered in Europe.5 Identification of evolutionary significant units and their management are crucial for conservation,6 especially as the white shark is facing various but often region-specific anthropogenic threats.7,8,9,10,11 Assessing connectivity in a cosmopolitan marine species requires worldwide sampling and high-resolution genetic markers.12 Both are lacking for the white shark, with studies to date typified by numerous but geographically limited sampling, and analyses relying largely on relatively small numbers of nuclear microsatellites,13,14,15,16,17,18,19 which can be plagued by various genotyping artefacts and thus require cautious interpretation.20 Sequencing and computational advances are finally allowing genomes21,22,23 to be leveraged into population studies,24,25,26,27 with datasets comprising thousands of single-nucleotide polymorphisms (SNPs). Here, combining target gene capture (TGC)28 sequencing (89 individuals, 4,000 SNPs) and whole-genome re-sequencing (17 individuals, 391,000 SNPs) with worldwide sampling across most of the distributional range, we identify three genetically distinct allopatric lineages (North Atlantic, Indo-Pacific, and North Pacific). These diverged 100,000-200,000 years ago during the Penultimate Glaciation, when low sea levels, different ocean currents, and water temperatures produced significant biogeographic barriers. Our results show that without high-resolution genomic analyses of samples representative of a species' range,12 the true extent of diversity, presence of past and contemporary barriers to gene flow, subsequent speciation, and local evolutionary events will remain enigmatic.

Adaptive spatiotemporal management to reduce shark bycatch in tuna fisheries.

Purse-seine tropical tuna fishing in the eastern tropical Pacific Ocean (EPO) results in the bycatch of several sensitive species groups, including elasmobranchs. Effective ecosystem management balances conservation and resource use and requires considering trade-offs and synergies. Seasonal and adaptive spatial measures can reduce fisheries impacts on nontarget species while maintaining or increasing target catches. Identifying persistently high-risk areas in the open ocean, where dynamic environmental conditions drive changes in species' distributions, is essential for exploring the impact of fisheries closures. We used fisheries observer data collected from 1995 to 2021 to explore the spatiotemporal persistence of areas of high bycatch risk for 2 species of oceanic sharks, silky shark (Carcharhinus falciformis) and oceanic whitetip shark (Carcharhinus longimanus), and of low tuna catch rates. We analyzed data collected by fisheries scientific observers onboard approximately 200 large purse-seine vessels operating in the EPO under 10 different flags. Fishing effort, catch, and bycatch data were aggregated spatially and temporally at 1° × 1° cells and monthly, respectively. When areas of high fishing inefficiency were closed the entire study period and effort was reallocated proportionally to reflect historical effort patterns, yearly tuna catch appeared to increase by 1-11%, whereas bycatch of silky and oceanic whitetip sharks decreased by 10-19% and 9%, respectively. Prior to fishing effort redistribution, bycatch reductions accrued to 21-41% and 14% for silky and oceanic whitetip sharks, respectively. Our results are consistent with previous findings and demonstrate the high potential for reducing elasmobranch bycatch in the EPO without compromising catch rates of target tuna species. They also highlight the need to consider new dynamic and adaptive management measures to more efficiently fulfill conservation and sustainability objectives for exploited resources in the EPO.

Gestión espaciotemporal adaptativa para reducir la captura incidental de tiburones en la pesca del atún Resumen La pesca con cerco del atún tropical en el Pacífico Tropical Oriental (PTO) resulta en la captura incidental de varios grupos de especies sensibles, incluidos los elasmobranquios. La gestión eficiente del ecosistema equilibra la conservación y el uso de recursos y requiere que se consideren las compensaciones y las sinergias. Las medidas espaciales adaptativas y estacionales pueden reducir el impacto de las pesquerías sobre las especies accesorias mientras mantienen o incrementan la captura intencional. La identificación de las áreas con alto riesgo persistente en mar abierto, en donde las condiciones ambientales dinámicas causan cambios en la distribución de las especies, es esencial para explorar el impacto del cierre de las pesquerías. Usamos datos de observadores de las pesquerías recolectados entre 1995 y 2021 para explorar la persistencia espaciotemporal de las áreas con alto riesgo de captura incidental para dos especies de tiburón (Carcharhinus falciformi y C. longimanus) y con tasas bajas de captura de atún. Analizamos los datos recolectados por los observadores científicos de las pesquerías a bordo de aproximadamente 200 embarcaciones grandes de pesca con cerco que operaban en el PTO bajo diez banderas diferentes. Agregamos los datos sobre el esfuerzo de pesca, captura y la captura incidental de forma espacial y temporal en celdas de 1° x 1° y mensual, respectivamente. Cuando las áreas con gran ineficiencia pesquera se encontraban cerradas durante toda la investigación y el esfuerzo se reasignaba proporcionalmente para reflejar los patrones históricos de esfuerzo, el esfuerzo anual de captura de atún parecía incrementar en un 1­11%, mientras que la captura incidental de las dos especies de tiburones disminuía en un 10­19% (C. falciformi) y 9% (C. longimanus). Antes de que de redistribuyera el esfuerzo de pesca, la reducción de la captura incidental se acumuló hasta el 21­41% (C. falciformi) y 14% (C. longimanus). Nuestros resultados son congruentes con resultados previos y demuestran el gran potencial de reducción de la captura incidental de elasmobranquios en el PTO sin poner en peligro las tasas de captura de las especies de atún. Los resultados también enfatizan la necesidad de considerar medidas adaptativas nuevas y dinámicas para cumplir de forma más eficiente los objetivos de conservación y sustentabilidad para la explotación de recursos en el PTO.

The effect of shortfin mako shark skin at the reattachment of a separated turbulent boundary layer.

This smooth flat experimental study investigates the capability of mako shark scales to control flow separation when placed downstream of the onset of turbulent boundary layer separation and within the reattachment region. The objective of the study is to validate the hypothesis that the shark scales' bristling and recoiling would prevent the flow separation on the flank region (the fastest flow region) of the shark. A rotating cylinder was used to induce an adverse pressure gradient over a flat plate to produce a region of separated flow where the shark skin specimen was mounted. Two types of mako shark scales (flank (B2) and between flank and dorsal fin (B1)) were positioned in the preferred flow direction on a flat plate. The B2 scales are slender, 200µm tall, and can bristle up to 50°. In contrast, B1 scales are wider, shorter, and can bristle at 30°. The bristling angle and shape are the main mechanisms by which the scales act to inhibit flow from moving upstream near the wall. Thus, the difference in the bristling angles and structures of the scales is attributed to the fact that the B2 scales function in a thicker boundary layer (behind the shark's gills) where they must bristle sufficiently high into the boundary layer to control the flow separation, and because the adverse pressure gradient in this region is higher where flow separation is more likely. The scales are placed in the reattachment region to elucidate their ability to control and reattach an already separated turbulent flow. The results show that B2 scales placed in the reattachment region reduce the size of the turbulent separation bubble and decrease the turbulent kinetic energy, while B1 scales have the opposite effect.

First report of recurrent parthenogenesis as an adaptive reproductive strategy in the endangered common smooth-hound shark Mustelus mustelus.

Parthenogenesis, or virgin birth, describes a mode of reproduction where an egg develops into an offspring without fertilization, and is observed across various vertebrate taxa, excluding mammals. Obligate parthenogenesis, found in around 100 vertebrate species and 1000 invertebrate species, is relatively rare. Conversely, facultative parthenogenesis, where females can reproduce both sexually and parthenogenetically, is observed in some vertebrates, including elasmobranchs. Notably, this phenomenon in elasmobranchs is mainly documented in captivity, allowing for detailed long-term observation. Specifically, this study reports the first case of facultative parthenogenesis in the common smooth-hound shark Mustelus mustelus, a species classified by IUCN as endangered. Here we show that the juvenile M. mustelus were born through parthenogenesis, exhibiting homozygosity at each genetic marker, consistent with terminal fusion automixis. Remarkably, this finding reveals that parthenogenesis can occur annually in these sharks, alternating between two females, and conclusively excludes long-term sperm storage as a cause. Consequently, this enhances our understanding of parthenogenesis in elasmobranchs and highlights the reproductive flexibility of M. mustelus. Overall, these results contribute to our broader understanding of reproductive strategies in elasmobranchs, which could inform conservation efforts for endangered species.

Laser Interference Additive Manufacturing: Mask Bundle Shape Bionic Shark Skin Structure.

Here, we explored a new manufacturing strategy that uses the mask laser interference additive manufacturing (MLIAM) technique, which combines the respective strengths of laser interference lithography and mask lithography to efficiently fabricate across-scales three-dimensional bionic shark skin structures with superhydrophobicity and adhesive reduction. The phenomena and mechanisms of the MLIAM curing process were revealed and analyzed, showing the feasibility and flexibility. In terms of structural performance, the adhesive force on the surface can be tuned based on the growth direction of the bionic shark skin structures, where the maximum rate of the adhesive reduction reaches about 65%. Furthermore, the evolution of the directional diffusion for the water droplet, which is based on the change of the contact angle, was clearly observed, and the mechanism was also discussed by the models. Moreover, no-loss transportations were achieved successfully using the gradient adhesive force and superhydrophobicity on the surface by tuning the growth direction and modifying by fluorinated silane. Finally, this work gives a strategy for fabricating across-scale structures on micro- and nanometers, which have potential application in bioengineering, diversional targeting, and condenser surface.

Characterization of shark skin properties and biomimetic replication.

This review explores the present knowledge of the unique properties of shark skin and possible applications of its functionalities, including drag reduction and swimming efficiency. Tooth-like denticles, with varied morphologies, sizes, and densities across the shark's body, significantly influence the flow and interaction of fluids. Examining dermal denticle morphology, this study unveils the functional properties of real shark skin, including mechanical properties such as stiffness, stress-strain characteristics, and denticle density's impact on tensile properties. The adaptive capabilities of the Mako shark scales, especially in high-speed swimming, are explored, emphasizing their passive flow-actuated dynamic micro-roughness. This research contains an overview of various studies on real shark skin, categorizing them into skin properties, morphology, and hydrodynamics. The paper extends exploration into industrial applications, detailing fabrication techniques and potential uses in vessels, aircraft, and water pipes for friction reduction. Three manufacturing approaches, bio-replicated forming, direct fabrication, and indirect manufacturing, are examined, with 3D printing and photoconfiguration technology emerging as promising alternatives. Investigations into the mechanical properties of shark skin fabrics reveal the impact of denticle size on tensile strength, stress, and strain. Beyond drag reduction, the study highlights the shark skin's role in enhancing thrust and lift during locomotion. The paper identifies future research directions, emphasizing live shark testing and developing synthetic skin with the help of 3D printing incorporating the bristling effect.

Platybothrium yanae sp. nov. (Cestoda: Onchobothriidae) from the whitecheek shark, Carcharhinus dussumieri (Valenciennes), off southern Iran.

Platybothrium Linton, 1890 is a genus parasitizing sharks of the families Carcharhinidae and Sphyrnidae. No new species has been assigned to the genus in the 20 years since its last treatment. In the present study, a new species is described from the Persian Gulf, which is the second report of a species of Platybothrium in the Indian Ocean. Platybothrium yanae sp. nov. differs from P. auriculatum Yamaguti, 1952, P. cervinum Linton, 1890, P. tantulum Healy, 2003, and P. kirstenae Healy, 2003 in lacking, rather than having, an accessory piece between its hooks. This new species is distinguished from its other congeners by having a particular combination of features including its measurements, morphology, and meristic features, bringing the number of valid species in the genus to 11.

Revealing Spermatogenesis in Smooth-Hound Sharks Mustelus mustelus: Insights into the Morphological and Macromolecular Composition of Spermatogenic Cells.

Elasmobranchs have an ancestral reproductive system, which offers insights into vertebrate reproductive evolution. Despite their unchanged design over 400 million years, they evolved complex mechanisms ensuring reproductive success. However, human activities induced a significant decline in elasmobranch populations worldwide. In the Mediterranean basin, the smooth-hound shark (Mustelus mustelus) is one of the species that are considered vulnerable to human activities. Conservation efforts necessitate a thorough understanding of its reproductive strategy. This study focused on mature male specimens of smooth-hound sharks that were captured in the Adriatic area and successively analyzed to provide, for the first time, a histologically detailed description of testicular development in the species. Seven phases of the spermatogenesis process were identified, along with the macromolecular characterization of cells obtained using Fourier-transform infrared imaging. Histological analysis showed structural and cellular features similar to those documented in the spermatocysts of other elasmobranchs. The examination of the evolution and migration of both germinative and Sertoli cells at each phase revealed their close connection. Furthermore, different expression levels of lipids, proteins, and phosphates (DNA) at each spermatogenesis stage were observed. This research provided new information on spermatogenesis in the common smooth-hound shark, which is crucial for conservation efforts against population decline and anthropogenic pressures.

The rise of pelagic sharks and adaptive evolution of pectoral fin morphology during the Cretaceous.

The emergence and subsequent evolution of pectoral fins is a key point in vertebrate evolution, as pectoral fins are dominant control surfaces for locomotion in extant fishes.1,2,3 However, major gaps remain in our understanding of the diversity and evolution of pectoral fins among cartilaginous fishes (Chondrichthyes), a group with an evolutionary history spanning over 400 million years with current selachians (modern sharks) appearing about 200 million years ago.4,5,6 Modern sharks are a charismatic group of vertebrates often thought to be predators roaming the open ocean and coastal areas, but most extant species occupy the seafloor.4 Here we use an integrative approach to understand what facilitated the expansion to the pelagic realm and what morphological changes accompanied this shift. On the basis of comparative analyses in the framework of a time-calibrated molecular phylogeny,7 we show that modern sharks expanded to the pelagic realm no later than the Early Cretaceous (Barremian). The pattern of pectoral fin aspect ratios across selachians is congruent with adaptive evolution, and we identify an increase of the subclade disparity of aspect ratio at a time when sea surface temperatures were at their highest.8 The expansion to open ocean habitats likely involved extended bouts of sustained fast swimming, which led to the selection for efficient movement via higher aspect ratio pectoral fins. Swimming performance was likely enhanced in pelagic sharks during this time due to the elevated temperatures in the sea, highlighting that shark evolution has been greatly impacted by climate change.

Maternal investment evolves with larger body size and higher diversification rate in sharks and rays.

Across vertebrates, live bearing evolved at least 150 times from ancestral egg laying into diverse forms and degrees of prepartum maternal investment.1,2 A key question is how reproductive diversity arose and whether reproductive diversification underlies species diversification.3,4,5,6,7,8,9,10,11 To test this, we evaluate the most basal jawed vertebrates: the sharks, rays, and chimaeras, which have one of the greatest ranges of reproductive and ecological diversity among vertebrates.2,12 We reconstruct the sequence of reproductive mode evolution across a phylogeny of 610 chondrichthyans.13 We reveal egg laying as ancestral, with live bearing evolving at least seven times. Matrotrophy evolved at least 15 times, with evidence of one reversal. In sharks, transitions to live bearing and matrotrophy are more prevalent in larger-bodied tropical species. Further, the evolution of live bearing is associated with a near doubling of the diversification rate, but there is only a small increase associated with the appearance of matrotrophy. Although pre-copulatory sexual selection is associated with increased rates of speciation in teleosts,3 sexual size dimorphism in chondrichthyans does not appear to be related to sexual selection,14,15 and instead we find increased rates of speciation associated with the colonization of novel habitats. This highlights a potential key difference between chondrichthyans and other fishes, specifically a slower rate of evolution of reproductive isolation following speciation, suggesting different rate-limiting mechanisms for diversification between these clades.16 The chondrichthyan diversification and radiation, particularly throughout shallow tropical shelf seas and oceanic pelagic habitats, appear to be associated with the evolution of live bearing and proliferation of a wide range of maternal investment in developing offspring.

The appendicular skeleton of the enigmatic shark Leptocharias smithii in comparison with other sharks of the order Carcharhiniformes (Elasmobranchii: Leptochariidae).

Leptocharias smithii has been poorly explored in anatomical terms. This species bears a mosaic of morphological characters and is considered to represent an intermediate condition between other carcharhiniform clades. In the present paper, the anatomy of the appendicular skeleton of the species is thoroughly investigated and compared with other representatives of the order Carcharhiniformes. Leptocharias bears exclusive characteristics, such as the visible separation of the pro- and mesopterygia but it also has an aplesodic pectoral fin, a condition shared with carcharhiniforms placed at the base of the phylogenetic tree and at the same time a chevron-shaped coracoid bar, a condition characteristic of charcharhiniforms placed at the apex of the phylogenetic tree. Additionally, in an attempt to understand the evolution of its appendicular skeleton and of other carcharhiniforms, 20 characters of the paired fins and girdles are explored and discussed in light of two recent phylogenetic hypotheses. Most of these characters were not previously explored and support not only the monophyly of Carcharhiniformes, such as the mesopterygium overlapping the metapterygium in ventral view, but also the monophyly of the less inclusive clade Hemigaleidae + (Galeocerdonidae + (Carcharhinidae+Sphyrnidae)), such as the morphology and arrangement of the distal radials, which are pointed and spaced.

Aspects of spermatogenesis in immature and mature specimens of the long-lived Greenland shark: Novelties concerning the germinal compartment's assembly, complement of Sertoli cells and demise.

Cystic spermatogenesis in the subadult, maturing and adult Greenland shark (Somniosus microcephalus) displays multiple novel features, characterized early on by an unorganized internal cellular environment of the spermatocysts (anatomically discrete follicle-like units containing a single germ cell stage and its complement of co-developing Sertoli cells). These typically show polar asymmetries due to asymmetrically distributed germ and Sertoli cells. These arise from several novel cellular rearrangements at the immature pole, including fusion of a cluster of somatic cells with newly formed cysts containing only one to three spermatogonia and that already display an excess of Sertoli cells. The subadult's germinative zone revealed an additional novelty, namely numerous previously formed somatic cell-lined rings into which spermatogonia were incorporated. A striking finding was the conspicuous rarity of the routinely discernible Sertoli mitotic figures in the hallmark cyst stage of diametric elasmobranch spermatogenesis that is known for the peak display of the latter. Scrutiny of sequentially unfolding phenomena in the linearly arranged spermatogonial generations revealed that the cellular developments at the most common type of cyst-duct transition area (comprising slender to spindle-like basophilic cells with pointed ends) were concurrent with the discreet appearance of a second dark Sertoli nucleus, a development that persisted in spermiated cysts. Spermatogenically active mature males displayed vigorous meiotic divisions. However, a scattering of their spermatid cysts also displayed shark-atypical asynchronous passage through spermiogenesis, phenomena which were exacerbated as arrested spermiogenesis in an archival collection of tissues from 13 maturing specimens. Subadult specimens revealed meiotic arrest, and foci of infiltration of leukocytes that originate from a mass of eosinophilic, granule-laden immune cells dorsally under the testis capsule. This tissue was identical to the testis-affixed bone marrow equivalent in other shark species. This tissue is likely developmentally regulated in the Greenland shark as it is absent in adults.