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.

Etmopterus lantern sharks use coelenterazine as the substrate for their luciferin-luciferase bioluminescence system.

The lantern shark genus Etmopterus contains approximately 40 species of deep-sea bioluminescent cartilaginous fishes. They emit blue light mainly from the ventral body surface. The biological functions of this bioluminescence have been discussed based on the luminescence patterns, but the bioluminescence mechanism remains uncertain. In this study, we detected both coelenterazine and coelenterazine-dependent luciferase activity in the ventral photophore tissue of Etmopterus molleri. The results suggested that bioluminescence in lantern sharks is produced using coelenterazine as the substrate for the luciferin-luciferase reaction, as some luminous bony fishes.

Evaluating artisanal fishing of globally threatened sharks and rays in the Bay of Bengal, Bangladesh.

Sharks and rays are at risk of extinction globally. This reflects low resilience to increasing fishing pressure, exacerbated by habitat loss, climate change, increasing value in a trade and inadequate information leading to limited conservation actions. Artisanal fisheries in the Bay of Bengal of Bangladesh contribute to the high levels of global fishing pressure on elasmobranchs. However, it is one of the most data-poor regions of the world, and the diversity, occurrence and conservation needs of elasmobranchs in this region have not been adequately assessed. This study evaluated elasmobranch diversity, species composition, catch and trade within the artisanal fisheries to address this critical knowledge gap. Findings show that elasmobranch diversity in Bangladesh has previously been underestimated. In this study, over 160000 individual elasmobranchs were recorded through landing site monitoring, comprising 88 species (30 sharks and 58 rays) within 20 families and 35 genera. Of these, 54 are globally threatened according to the IUCN Red List of Threatened Species, with ten species listed as Critically Endangered and 22 species listed as Endangered. Almost 98% juvenile catch (69-99% for different species) for large species sand a decline in numbers of large individuals were documented, indicating unsustainable fisheries. Several previously common species were rarely landed, indicating potential population declines. The catch pattern showed seasonality and, in some cases, gear specificity. Overall, Bangladesh was found to be a significant contributor to shark and ray catches and trade in the Bay of Bengal region. Effective monitoring was not observed at the landing sites or processing centres, despite 29 species of elasmobranchs being protected by law, many of which were frequently landed. On this basis, a series of recommendations were provided for improving the conservation status of the elasmobranchs in this region. These include the need for improved taxonomic research, enhanced monitoring of elasmobranch stocks, and the highest protection level for threatened taxa. Alongside political will, enhancing national capacity to manage and rebuild elasmobranch stocks, coordinated regional management measures are essential.

International fisheries threaten globally endangered sharks in the Eastern Tropical Pacific Ocean: the case of the Fu Yuan Yu Leng 999 reefer vessel seized within the Galápagos Marine Reserve.

Shark fishing, driven by the fin trade, is the primary cause of global shark population declines. Here, we present a case study that exemplifies how industrial fisheries are likely depleting shark populations in the Eastern Tropical Pacific Ocean. In August 2017, the vessel Fu Yuan Yu Leng 999, of Chinese flag, was detained while crossing through the Galápagos Marine Reserve without authorization. This vessel contained 7639 sharks, representing one of the largest seizures recorded to date. Based on a sample of 929 individuals (12%), we found 12 shark species: 9 considered as Vulnerable or higher risk by the IUCN and 8 listed in CITES. Four species showed a higher proportion of immature than mature individuals, whereas size-distribution hints that at least some of the fishing ships associated with the operation may have been using purse-seine gear fishing equipment, which, for some species, goes against international conventions. Our data expose the magnitude of the threat that fishing industries and illegal trade represent to sharks in the Eastern Tropical Pacific Ocean.

Fossil dermal denticles reveal the preexploitation baseline of a Caribbean coral reef shark community.

Preexploitation shark baselines and the history of human impact on coral reef-associated shark communities in the Caribbean are tpoorly understood. We recovered shark dermal denticles from mid-Holocene (∼7 ky ago) and modern reef sediments in Bocas del Toro, Caribbean Panama, to reconstruct an empirical shark baseline before major human impact and to quantify how much the modern shark community in the region had shifted from this historical reference point. We found that denticle accumulation rates, a proxy for shark abundance, declined by 71% since the mid-Holocene. All denticle morphotypes, which reflect shark community composition, experienced significant losses, but those morphotypes found on fast-swimming, pelagic sharks (e.g., families Carcharhinidae and Sphyrnidae) declined the most. An analysis of historical records suggested that the steepest decline in shark abundance occurred in the late 20th century, coinciding with the advent of a targeted shark fishery in Panama. Although the disproportionate loss of denticles characterizing pelagic sharks was consistent with overfishing, the large reduction in denticles characterizing demersal species with low commercial value (i.e., the nurse shark Ginglymostoma cirratum) indicated that other stressors could have exacerbated these declines. We demonstrate that the denticle record can reveal changes in shark communities over long ecological timescales, helping to contextualize contemporary abundances and inform shark management and ecology.

Identifying shark species responsible for fisheries depredation off Southeast Queensland, Australia.

Anecdotal reports from fishers in Southeast Queensland, Australia suggest that shark depredation is a significant issue, however little is known about which species are responsible for depredating catches. This research aimed to identify depredating species in Southeast Queensland line based fisheries, by undertaking a genetic analysis of depredated samples collected by commercial, charter and recreational fishers. The genetic analysis successfully identified ten depredating sharks, all from the genus Carcharhinus (19.2% success). The species identified using mitochondrial DNA included five C. leucas (bull sharks), two C. plumbeus (sandbar sharks), one C. amboinensis (pigeye shark), one C. brevipinna (spinner shark) and one unconfirmed C. plumbeus/C. altimus (bignose shark). While many species of Carcharhinus have been found to depredate catches in Australia, C. leucas has not been highlighted until this research as a potential problematic species. The optimised protocol allowed for the confident identification of shark species responsible for depredation in fisheries using frozen fish samples donated by fishers.

Manta-like planktivorous sharks in Late Cretaceous oceans.

The ecomorphological diversity of extinct elasmobranchs is incompletely known. Here, we describe Aquilolamna milarcae, a bizarre probable planktivorous shark from early Late Cretaceous open marine deposits in Mexico. Aquilolamna, tentatively assigned to Lamniformes, is characterized by hypertrophied, slender pectoral fins. This previously unknown body plan represents an unexpected evolutionary experimentation with underwater flight among sharks, more than 30 million years before the rise of manta and devil rays (Mobulidae), and shows that winglike pectoral fins have evolved independently in two distantly related clades of filter-feeding elasmobranchs. This newly described group of highly specialized long-winged sharks (Aquilolamnidae) displays an aquilopelagic-like ecomorphotype and may have occupied, in late Mesozoic seas, the ecological niche filled by mobulids and other batoids after the Cretaceous-Paleogene boundary.

Complete Mitochondrial DNA Genome of Nine Species of Sharks and Rays and Their Phylogenetic Placement among Modern Elasmobranchs.

Chondrichthyes occupy a key position in the phylogeny of vertebrates. The complete sequence of the mitochondrial genome (mitogenome) of four species of sharks and five species of rays was obtained by whole genome sequencing (DNA-seq) in the Illumina HiSeq2500 platform. The arrangement and features of the genes in the assembled mitogenomes were identical to those found in vertebrates. Both Maximum Likelihood (ML) and Bayesian Inference (BI) analyses were used to reconstruct the phylogenetic relationships among 172 species (including 163 mitogenomes retrieved from GenBank) based on the concatenated dataset of 13 individual protein coding genes. Both ML and BI analyses did not support the "Hypnosqualea" hypothesis and confirmed the monophyly of sharks and rays. The broad notion in shark phylogeny, namely the division of sharks into Galeomorphii and Squalomorphii and the monophyly of the eight shark orders, was also supported. The phylogenetic placement of all nine species sequenced in this study produced high statistical support values. The present study expands our knowledge on the systematics, genetic differentiation, and conservation genetics of the species studied, and contributes to our understanding of the evolutionary history of Chondrichthyes.

Paracellular and Transcellular Leukocytes Diapedesis Are Divergent but Interconnected Evolutionary Events.

Infiltration of the endothelial layer of the blood-brain barrier by leukocytes plays a critical role in health and disease. When passing through the endothelial layer during the diapedesis process lymphocytes can either follow a paracellular route or a transcellular one. There is a debate whether these two processes constitute one mechanism, or they form two evolutionary distinct migration pathways. We used artificial intelligence, phylogenetic analysis, HH search, ancestor sequence reconstruction to investigate further this intriguing question. We found that the two systems share several ancient components, such as RhoA protein that plays a critical role in controlling actin movement in both mechanisms. However, some of the key components differ between these two transmigration processes. CAV1 genes emerged during Trichoplax adhaerens, and it was only reported in transcellular process. Paracellular process is dependent on PECAM1. PECAM1 emerged from FASL5 during Zebrafish divergence. Lastly, both systems employ late divergent genes such as ICAM1 and VECAM1. Taken together, our results suggest that these two systems constitute two different mechanical sensing mechanisms of immune cell infiltrations of the brain, yet these two systems are connected. We postulate that the mechanical properties of the cellular polarity is the main driving force determining the migration pathway. Our analysis indicates that both systems coevolved with immune cells, evolving to a higher level of complexity in association with the evolution of the immune system.

A method for making alignments of related protein sequences that share very little similarity; shark interleukin 2 as an example.

An optimized alignment of related protein sequences helps to see their important shared features and to deduce their phylogenetic relationships. At low levels of sequence similarity, there are no suitable computer programs for making the best possible alignment. This review summarizes some guidelines for how in such instances, nevertheless, insightful alignments can be made. The method involves, basically, the understanding of molecular family features at both the protein and intron-exon level, and the collection of many related sequences so that gradual differences may be observed. The method is exemplified by identifying and aligning interleukin 2 (IL-2) and related sequences in Elasmobranchii (sharks/rays) and coelacanth, as other authors have expressed difficulty with their identification. From the point of general immunology, it is interesting that the unusual long "leader" sequence of IL-15, already known in other species, is even more impressively conserved in cartilaginous fish. Furthermore, sequence comparisons suggest that IL-2 in cartilaginous fish has lost its ability to bind an IL-2Rα/15Rα receptor chain, which would prohibit the existence of a mechanism for regulatory T cell regulation identical to mammals.

Coastal sharks and rays in a marine protected area in southern Brazil.

In southern Brazil, we investigated shark and ray capture records in an integral protection marine protection area (MPA) that allows fishing in that area. We found 10 shark and nine ray species, of which 88.5% and 66.2%, respectively, are endangered. Female adults and neonates of both sexes are abundant in the area, with records of stranding of large specimens, postcapture abortions and ovigerous capsules with fresh embryos on the beach. We suggest the application of continuous fisheries monitoring and an increase in guidelines on the capture of endangered species.

Isotopic (δ15 N) relationship of pregnant females and their embryos: Comparing placental and yolk-sac viviparous elasmobranchs.

Nitrogen stable isotopes ratios (δ15 N) were determined for selected tissues (muscle, liver, blood and yolk) of pregnant females and their embryos of a placental viviparous species, the Pacific sharpnose shark (Rhizoprionodon longurio), and a yolk-sac viviparous species, the speckled guitarfish (Pseudobatos glaucostigmus). The R. longurio embryo tissues were 15 N enriched compared to the same tissues in the pregnant female, using the difference in δ15 N (Δδ15 N) between embryo and adult. Mean Δδ15 N was 2.17‰ in muscle, 4.39‰ in liver and 0.80‰ in blood. For P. glaucostigmus, embryo liver tissue was significantly 15 N enriched in comparison with liver of the pregnant female (Δδ15 N mean = 1.22‰), whereas embryo muscle was 15 N depleted relative to the muscle of the pregnant female (Δδ15 N mean = -1.22‰). Both species presented a significant positive linear relationship between Δδ15 N and embryo total length (LT ). The results indicated that embryos have different Δδ15 N depending on their reproductive strategy, tissue type analysed and embryo LT .

A reappraisal of the pectoral skeleton of lantern sharks (Elasmobranchii: Squaliformes: Etmopteridae).

The morphology of the articular region of the pectoral girdle and associated basals in Etmopteridae is revised in light of new evidence provided by taxa unavailable for previous studies. Such studies considered that etmopterids plesiomorphically had a single pectoral articular condyle, and only Etmopterus had two separate ones. Our reanalysis indicates that the possession of two separate condyles, one for the articulation of the propterygium and the second for the meso- and metapterygium, is the most widespread condition in this group. However, the presence of two separate articular condyles is not recovered as a synapomorphy for Etmopteridae. Previous studies also proposed that etmopterids lack a hook-like process on the anteroproximal margin of the anteriormost pectoral basal. We document that the hook-like process is plesiomorphically present in Etmopteridae, thus corroborating the hypothesis of closer relationships between this family and the other squaliforms that also share this process, namely Centrophoridae, Dalatiidae, Oxynotidae, and Somniosidae.

Architectural and ultrastructural features of tessellated calcified cartilage in modern and extinct chondrichthyan fishes.

Tessellated calcified cartilage (TCC) is a distinctive kind of biomineralized perichondral tissue found in many modern and extinct chondrichthyans (sharks, rays, chimaeroids and their extinct allies). Customarily, this feature has been treated somewhat superficially in phylogenetic analyses, often as a single "defining" character of a chondrichthyan clade. TCC is actually a complex hard tissue with numerous distinctive attributes, but its use as a character complex for phylogenetic analysis has not yet been optimized. This study attempts to improve this situation by presenting new terminology for certain aspects of tesseral architecture, including single-monolayered, multiple-monolayered, polylayered and voussoir tesserae; new histological data, including thin sections of TCC in several Palaeozoic taxa, and new proposals for ways in which various characters and states (many of which are defined here for the first time) could be applied in future phylogenetic analyses of chondrichthyan fishes. It can be concluded that many, but not all, of the unique attributes of modern TCC evolved by the Early Devonian (ca. 400 before present (bp)). The globular calcified cartilage reported in Silurian sinacanthids and the so-called subtessellated perichondral biomineralization (with irregular and ill-defined geometries of a layer or layers of calcified cartilage blocks) of certain extinct "acanthodians" (e.g., Climatius, Ischnacanthus, Cheiracanthus) could represent evolutionary precursors of TCC, which seems to characterize only part of the chondrichthyan total group. It is hypothesized that heavily biomineralized "layer-cake" TCC in certain Palaeozoic chondrichthyans perhaps served a dual physiological role, as a phosphate sink and in providing increased skeletal density in very large (>7 m) Devonian-Permian marine sharks such as ctenacanths and as an adaptation to calcium-deficient environments among Permo-Carboniferous non-marine sharks such as xenacanths. By contrast, the equivalent tissue in modern elasmobranchs probably serves only to reinforce regions of cartilage (mostly in the jaws) subjected to high loading. It is also noted that much of the variation observed in tesseral architecture (including localized remodelling), ultrastructure and histology in modern and extinct chondrichthyans is confined to the perichondrally facing cap zone (where Type-1 collagen matrix predominates in modern TCC), whereas the main body of the tessera (where Type-2 collagen matrix predominates) exhibits comparatively little evidence of remodelling and histological or structural variation.

Catch composition and life history characteristics of sharks and rays (Elasmobranchii) landed in the Andaman and Nicobar Islands, India.

Detailed information on shark and ray fisheries in the Andaman and Nicobar Islands, India are limited, including information on the diversity and biological characteristics of these species. We carried out fish landing surveys in South Andamans from January 2017 to May 2018, a comprehensive and cost-effective way to fill this data gap. We sampled 5,742 individuals representing 57 shark and ray species landed from six types of fishing gears. Of the 36 species of sharks and 21 species of rays landed, six species of sharks (Loxodon macrorhinus, Carcharhinus amblyrhynchos, Sphyrna lewini, C. albimarginatus, C. brevipinna, and Paragaleus randalli) comprised 83.35% of shark landings, while three species of rays (Pateobatis jenkinsii, Himantura leoparda and H. tutul) comprised 48.82% of ray landings, suggesting a species dominance in the catch or fishing region. We provide insights into the biology of species with extensions in maximum size for seven shark species. Additionally, we document an increase in the known ray diversity for the islands and for India with three previously unreported ray species. We found that amongst sharks, mature individuals of small-bodied species (63.48% males of total landings of species less than 1.5 m total length when mature) and immature individuals of larger species (84.79% males of total landings of species larger than 1.5 m total length when mature) were mostly landed; whereas for rays, mature individuals were predominantly landed (80.71% males of total landings) likely reflecting differences in habitat preferences along life-history stages across species and fishing gear. The largest size range in sharks was recorded in landings from pelagic longlines and gillnets. Further, the study emphasizes the overlap between critical habitats and fishing grounds, where immature sharks and gravid females were landed in large quantities which might be unsustainable in the long-term. Landings were female-biased in C. amblyrhynchos, S. lewini and P. jenkinsii, and male-biased in L. macrorhinus and H. leoparda, indicating either spatio-temporal or gear-specific sexual segregation in these species. Understanding seasonal and biological variability in the shark and ray landings over a longer study period across different fisheries will inform future conservation and fishery management measures for these species in the Andaman and Nicobar Islands.

Evolution of vertebrate gill covers via shifts in an ancient Pou3f3 enhancer.

Whereas the gill chambers of jawless vertebrates open directly into the environment, jawed vertebrates evolved skeletal appendages that drive oxygenated water unidirectionally over the gills. A major anatomical difference between the two jawed vertebrate lineages is the presence of a single large gill cover in bony fishes versus separate covers for each gill chamber in cartilaginous fishes. Here, we find that these divergent patterns correlate with the pharyngeal arch expression of Pou3f3 orthologs. We identify a deeply conserved Pou3f3 arch enhancer present in humans through sharks but undetectable in jawless fish. Minor differences between the bony and cartilaginous fish enhancers account for their restricted versus pan-arch expression patterns. In zebrafish, mutation of Pou3f3 or the conserved enhancer disrupts gill cover formation, whereas ectopic pan-arch Pou3f3b expression generates ectopic skeletal elements resembling the multimeric covers of cartilaginous fishes. Emergence of this Pou3f3 arch enhancer >430 Mya and subsequent modifications may thus have contributed to the acquisition and diversification of gill covers and respiratory strategies during gnathostome evolution.

Species composition of the largest shark fin retail-market in mainland China.

Species-specific monitoring through large shark fin market surveys has been a valuable data source to estimate global catches and international shark fin trade dynamics. Hong Kong and Guangzhou, mainland China, are the largest shark fin markets and consumption centers in the world. We used molecular identification protocols on randomly collected processed fin trimmings (n = 2000) and non-parametric species estimators to investigate the species composition of the Guangzhou retail market and compare the species diversity between the Guangzhou and Hong Kong shark fin retail markets. Species diversity was similar between both trade hubs with a small subset of species dominating the composition. The blue shark (Prionace glauca) was the most common species overall followed by the CITES-listed silky shark (Carcharhinus falciformis), scalloped hammerhead shark (Sphyrna lewini), smooth hammerhead shark (S. zygaena) and shortfin mako shark (Isurus oxyrinchus). Our results support previous indications of high connectivity between the shark fin markets of Hong Kong and mainland China and suggest that systematic studies of other fin trade hubs within Mainland China and stronger law-enforcement protocols and capacity building are needed.

Diafanización dental de cuatro especies de seláceos (Carcharhinus leucas, Galeocerdo cuvier, Rhizoprionodon longurio y Sphyrna sp) / Dental diaphonization of four salaceos species (Carcharhinus leucas, Galeocerdo Cuvier, Rhizoprionodon longurio and Sphyrna sp)

El estudio morfológico de la dentadura de chondrichthyes representa un carácter taxonómico importante empleado para la clasificación e identificación de diferentes especies. Se diafanizaron dientes de cuatro especies distintas de selacimorfos (Carcharhinus leucas, Galeocerdo cuvier, Rhizoprionodon longurio y Sphyrna sp.) con la finalidad de estandarizar una técnica dental para su transparentación. Estandarizando la técnica de Okumura-Aprile aplicada para la diafanización dental de humanos, se obtuvo una diafanización óptima en las cuatro especies en tratamiento con HCl al 7 % donde se podía observar con claridad la cámara pulpar, por lo que podemos concluir que la técnica de Okumura-Aprile es eficiente en la diafanización dental de tiburones.

The morphological study of the chondrichthyes teeth represents an important taxonomic characteristic used for the classification and identification of different species. The teeth of four different species of selacimorphs (Carcharhinus leucas, Galeocerdo cuvier, Rhizoprionodon longurio and Sphyrna sp.) were diaphonized in order to standardize a dental technique for their transparency. By standardizing the Okumura-Aprile technique applied for the dental diaphonization of humans, an optimal diaphonization was obtained in the four species treated with 7 % HCl where the pulp chamber was clearly observed. Therefore, we may conclude that the OkumuraAprile technique is efficient in shark dental diaphonization.

Sharks, rays, and chimaeras of the Revillagigedo National Park: An update of new and confirmed records.

The Revillagigedo Archipelago National Park is the largest no-take Marine Protected Area in North America and it is essential for the conservation of globally endangered marine megafauna, including cetaceans, sea birds and cartilaginous fishes. The present paper reports the occurrence of 28 chondrichthyan species, including 19 sharks, seven batoids and two chimaeras in the Revillagigedo Archipelago, comprising eight new records and a brief revision of confirmed species to date.

Discovery of a new mode of oviparous reproduction in sharks and its evolutionary implications.

Two modes of oviparity are known in cartilaginous fishes, (1) single oviparity where one egg case is retained in an oviduct for a short period and then deposited, quickly followed by another egg case, and (2) multiple oviparity where multiple egg cases are retained in an oviduct for a substantial period and deposited later when the embryo has developed to a large size in each case. Sarawak swellshark Cephaloscyllium sarawakensis of the family Scyliorhinidae from the South China Sea performs a new mode of oviparity, which is named "sustained single oviparity", characterized by a lengthy retention of a single egg case in an oviduct until the embryo attains a sizable length. The resulting fecundity of the Sarawak swellshark within a season is quite low, but this disadvantage is balanced by smaller body, larger neonates and quicker maturation. The Sarawak swellshark is further uniquely characterized by having glassy transparent egg cases, and this is correlated with a vivid polka-dot pattern of the embryos. Five modes of lecithotrophic (yolk-dependent) reproduction, i.e. short single oviparity, sustained single oviparity, multiple oviparity, yolk-sac viviparity of single pregnancy and yolk-sac viviparity of multiple pregnancy were discussed from an evolutionary point of view.