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.

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before major human impact and to quantify how much the modern shark community in the region had shifted from this historical reference point.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.

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.

An early chondrichthyan and the evolutionary assembly of a shark body plan.

Although relationships among the major groups of living gnathostomes are well established, the relatedness of early jawed vertebrates to modern clades is intensely debated. Here, we provide a new description of Gladbachus, a Middle Devonian (Givetian approx. 385-million-year-old) stem chondrichthyan from Germany, and one of the very few early chondrichthyans in which substantial portions of the endoskeleton are preserved. Tomographic and histological techniques reveal new details of the gill skeleton, hyoid arch and jaws, neurocranium, cartilage, scales and teeth. Despite many features resembling placoderm or osteichthyan conditions, phylogenetic analysis confirms Gladbachus as a stem chondrichthyan and corroborates hypotheses that all acanthodians are stem chondrichthyans. The unfamiliar character combination displayed by Gladbachus, alongside conditions observed in acanthodians, implies that pre-Devonian stem chondrichthyans are severely under-sampled and strongly supports indications from isolated scales that the gnathostome crown group originated at the latest by the early Silurian (approx. 440 Ma). Moreover, phylogenetic results highlight the likely convergent evolution of conventional chondrichthyan conditions among earliest members of this primary gnathostome division, while skeletal morphology points towards the likely suspension feeding habits of Gladbachus, suggesting a functional origin of the gill slit condition characteristic of the vast majority of living and fossil chondrichthyans.

Denticle-embedded ampullary organs in a Cretaceous shark provide unique insight into the evolution of elasmobranch electroreceptors.

Here, we report a novel type of dermal denticle (or placoid scale), unknown among both living and fossil chondrichthyan fishes, in a Cretaceous lamniform shark. By their morphology and location, these dermal denticles, grouped into clusters in the cephalic region, appear to have been directly associated with the electrosensory ampullary system. These denticles have a relatively enlarged (∼350 µm in diameter), ornamented crown with a small (∼100 µm) asterisk- or cross-shaped central perforation connected to a multi-alveolate internal cavity. The formation of such a complex structure can be explained by the annular coalescence and fusion, around an ampullary vesicle, of several developmental units still at papillary stage (i.e. before mineralization), leading to a single denticle embedding an alveolar ampulla devoid of canal. This differs from larger typical ampullae of Lorenzini with a well-developed canal opening in a pore of the skin and may represent another adaptive response to low skin resistance. Since it has been recently demonstrated that ampullary organs arise from lateral line placodes in chondrichthyans, this highly specialized type of dermal denticle (most likely non-deciduous) may be derived from the modified placoid scales covering the superficial neuromasts (pit organs) of the mechanosensory lateral line system of many modern sharks.

A new Xenacanthiformes shark (Chondrichthyes, Elasmobranchii) from the Late Paleozoic Rio do Rasto Formation (Paraná Basin), Southern Brazil.

The Brazilian records on Xenacanthiformes include teeth and cephalic spines from the Parnaíba, Amazonas and Paraná basins. This work describes a new species of Xenacanthidae, collected in an outcrop of Serrinha Member of Rio do Rasto Formation (Wordian to Wuchiapingian), Paraná Basin, municipality of Jacarezinho, State of Paraná. The teeth of the new species are two or three-cuspidated and the aboral surface show a smooth concavity and one rounded basal tubercle. The coronal surface presents one semi-spherical and subcircular coronal button, and also two lateral main cusps and one central (when present) with less than one fifth of the size of the lateral cusps in the labial portion. The lateral cusps are asymmetric or symmetric, rounded in transversal section, lanceolate in longitudinal section, devoid of lateral carinae and lateral serrations, and with few smooth cristae of enameloid. In optical microscope the teeth show a trabecular dentine (osteodentine) base, while the cusps are composed by orthodentine, and the pulp cavities are non-obliterated by trabecular dentine. The fossil assemblage in the same stratigraphical level and in the whole Rio do Rasto Formation indicates another freshwater record for xenacanthid sharks.

Redescription of Hexanchus nakamurai Teng 1962, (Chondrichthyes: Hexanchiformes: Hexanchidae), with designation of a neotype.

Hexanchus nakamurai Teng, 1962 is one of two valid nominal species in the genus Hexanchus; the other being H. griseus. The taxonomic history of H. nakamurai is somewhat convoluted due to questions about the validity of whether it constituted a publication or an unpublished dissertation. The issue appeared to have been resolved once it was determined that the Teng's original description met the criteria under the International Commission on Zoological Nomenclature code (Article 8a) of a publication. However, recent molecular studies have indicated that the western North Atlantic H. nakamurai (= ? H. vitulus) may be distinct from western Indian Ocean H. nakamurai. Compounding the issue is the loss of the holotype of H. nakamurai from Taiwan. A neotype is herein designated and the species redescribed based on the neotype and eight additional Taiwanese specimens.

Morphological and taxonomic revision of species of Squatina from the Southwestern Atlantic Ocean (Chondrichthyes: Squatiniformes: Squatinidae).

The morphology and taxonomy of species of Squatina from the southwestern Atlantic Ocean are revised. Species previously considered valid, Squatina argentina (Marini, 1930), Squatina guggenheim Marini, 1936 and Squatina occulta Vooren and da Silva, 1991, are investigated and described in detail, including a morphometric and meristic study of specimens from their recorded range. The taxonomic status of the doubtful nominal species Squatina punctata Marini, 1936 was also evaluated. This species was previously considered a junior synonym of S. argentina, a junior synonym of S. guggenheim, or a senior synonym of S. occulta. Although there is much morphological similarity between Squatina species, significant differences in dermal denticle patterns, dorsal coloration, tooth formula, and size at maturity are reported, enabling the recognition of S. argentina, S. guggenheim and S. occulta as valid species, and relegating S. punctata to the synonymy of S. guggenheim. Differences in skeletal morphology between valid species are described and considered supportive of the taxonomic hypothesis, corroborating a previous study of neurocrania. Additionally, an unidentified specimen is reported, as Squatina sp., from the continental shelf of Bahia state, Brazil, recognized by having more vertebral centra and a conspicuous dermal denticle morphology on interspiracular region, features not present in other South America angelshark species. A report on the only known syntype of Squatina dumeril Le Sueur, 1818 is presented, describing features that are still preserved and designating it as lectotype. Lateral-line sensory canals, skeleton, and cranial and hypobranchial muscles for the three valid species of Squatina from the southwestern Atlantic, as well as the brain and cranial nerves of S. guggenheim, are described and illustrated.

Revision of the genus Centrophorus (Squaliformes: Centrophoridae): Part 1--Redescription of Centrophorus granulosus (Bloch & Schneider), a senior synonym of C. acus Garman and C. niaukang Teng.

The genus Centrophorus is one of the most taxonomically complex and confusing elasmobranch groups. A revision of this group is currently underway and this first paper sets an important foundation in this process by redescribing the type species of the genus--Centrophorus granulosus. This taxon name has been previously applied to two different morphotypes: a large species > 1.5 m TL and a smaller species -1 m TL. Centrophorus acus and C. niaukang are the most commonly used names applied to the larger morphotype. The original description of C. granulosus was based on a large specimen of -1.5 m TL, but subsequent redescriptions were based on either of the large or small morphotypes. Centrophorus granulosus is herein redescribed as a large species and a neotype is designated. Centrophorus acus and C. niaukang are found to be junior synonyms of C. granulosus. Centrophorus granulosus is distinguishable from its congeners by its large size, dermal denticle shape, colouration and a number of morphological and biological characteristics. Ontogenetic changes in morphology, dentition and denticle shape for this species are described in detail.

Lost and found: recovery of the holotype of the ocellated angelshark, Squatina tergocellatoides Chen, 1963 (Squatinidae), with comments on western Pacific squatinids.

The ocellated angelshark, Squatina tergocellatoides, Chen, 1963 is redescribed from the holotype, which was thought to be lost. Its recent recovery has allowed for a revised description, including new data, and comparison to other Western Pacific squatinids. Squatina tergocellatoides can be distinguished from its congeners by three pairs of prominent large black spots, each with a diameter greater than eye length; two on each pectoral fin at anterior and posterior angles and one on each side near the tail base; another three pairs of lesser defined spots, one large spot on base of each dorsal fin and one located laterally on each side of tail located below first dorsal fin. Ventral surface is uniformly white to cream coloured, and margins of pectoral fins and tail similar in colour to dorsal side. Pectoral fins with angular lateral apices and rounded posterior lobe, pelvic fin tips not reaching origin of first dorsal fin, strongly fringed nasal barbels, small inter-orbital space, head and mouth lengths, broad internarial width and pelvic fin base, a very small pelvic girdle width, and a caudal fin with triangular ventral lobe greater in length than dorsal lobe. Comments on additional specimens are provided, as well as observations on biogeography. A review of western Pacific squatinids is also provided.

Pristiophorus lanae sp. nov., a new sawshark species from the western North Pacific, with comments on the genus Pristiophorus Müller & Henle, 1837 (Chondrichthyes: Pristiophoridae).

A new species of sawshark, Pristiophorus lanae sp. nov., is described from off the Philippine Islands. The new species is the second member of the genus Pristiophorus described from the western North Pacific and can be separated from its closest geographic congener, P. japonicus, by having fewer rostral teeth in front of rostral barbels (17-26 versus 25-32), mouth at corners extending forward to below the rear margin of the eye versus extending below the rear one-third of eye margin, a greater mouth width at 6.9-7.8 times into pre-oral length (versus 5.8-6.9), eye length into head length (15.6-15.9 versus 9.8-13.2), mouth width into head length 9.0-10.0 versus 7.4-8.5 times, head width at nostrils 5.2-6.1 times into pre-orbital length versus 3.9-4.9 times, shorter prebarbel length (from snout tip to barbel) of 50.7-54.5% of preoral length versus 53.6-59.2%, a snout angle of 10.6-13.0 degrees versus 12.4 degrees - 14.6 degrees, and lateral trunk denticles with flat crowns that are imbricated versus erect crowns that are not imbricated. The number of monospondylous vertebrae is slightly lowe in P. lanae (43-48) versus P. japonicus (51-52). The genus is reviewed, with a revised key to its species presented.

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.

The oldest articulated chondrichthyan from the Early Devonian period.

Chondrichthyans (including living sharks, skates, rays and chimaeras) have a fossil record of scales and dermal denticles perhaps dating back to the Late Ordovician period, about 455 million years ago. Their fossil tooth record extends to the earliest Devonian period, almost 418 million years ago, whereas the oldest known articulated shark remains date from the Early Devonian period, about 394 million years ago. Here we report the discovery of an articulated shark that is almost 409 million years old from the Early Devonian (early Emsian) period of New Brunswick, Canada. The specimen, identified as Doliodus problematicus (Woodward), sheds light on the earliest chondrichthyans and their interrelationships with basal jawed vertebrates. This species has been truly problematic. Previously known only from isolated teeth, it has been identified as an acanthodian and a chondrichthyan. This specimen is the oldest shark showing the tooth families in situ, and preserves one of the oldest chondrichthyan braincases. More notably, it shows the presence of paired pectoral fin-spines, previously unknown in cartilaginous fishes.

Redefining species concepts for the Pennsylvanian scissor tooth shark, Edestus.

This study reevaluates the tooth morphology used to define species within the genus Edestus (Chondrichthyes, Euchondrocephali). Known as the scissor tooth shark, Edestus produced a unique dentition of spiraled tooth families positioned in the symphysis (midline) of the upper and lower jaws. Morphometric analysis of more than 200 ejected teeth and intact spiral tooth whorls demonstrates that teeth from the upper and lower whorls differ in shape and ontogeny. Comparison of these data to the type specimens of 13 existing species reduces the number of morphologically distinct Edestus to just four species and refines the stratigraphic occurrence and expansion of the group. E. triserratus has a narrow bullet-shaped crown that points anteriorly and has roots of intermediate length. E. minor crowns have a wider triangular base, whereas the crowns of E. heinrichi form nearly equilateral triangles and are supported by an elongated root. E. vorax, which also has roughly equilateral triangular crowns, has short and deep roots, and is only known from very large specimens that are distinct from the growth series of E. heinrichi. Tooth and whorl morphologies among the species are consistent with cranial anatomy observed in a juvenile E. heinrichi and with transverse tooth-wear patterns to suggest Edestus used a forward to backward slicing motion to bite its prey. Extrapolating body size from tooth whorl length provides a conservative estimate that E. heinrichi could exceed 6.7 m in length. Edestus fossils are recovered from coastal marine to estuarine deposits spanning roughly six million years (313-307 Ma). Edestus first appears in England during the latest Bashkirian (313 Ma, Carboniferous), a few million years after its most closely resembling genus Lestrodus. Diversification and range expansion of Edestus coincides with the Moscovian transgression that flooded Laurentia and the Russian platform.

Shape analysis of shark jaws as a tool to identify species involved in incidents with humans.

Shark incidents in Recife, Pernambuco state (Brazil), are registered since 1992. The occurrence of those events is in a relatively small area of urban beaches of about 30 km long, making up one of the world's highest indices of shark incidents per unit of area, totaling 64 casualties, of these 24 were fatal. To mitigate shark incidents, the knowledge on shark species involved is crucial given that each species has specific usage of the habitat and dissimilar feeding strategy. Jaws of five species caught along the Pernambuco coast (Brazil), corresponded to 2 specimens of Carcharhinus acronotus, 4 specimens of C. leucas, 1 of C. limbatus, 4 of Galeocerdo curvier and 3 of Sphyrna lewini. Relative Warps Analysis was applied to enhance possible differences among jaw shapes of the studied species. The consensus configuration from the 14 images was obtained. The first two relative warps components explained 70.17% of the total observed variation. Four distinct groups were clearly observed. Three groups were formed by single species, corresponding to G. curvier, C. leucas and S. lewini, respectively, and one group formed by the combination of C. acronotus and C. limbatus. As expected, due to its peculiar skull shape, S. lewini jaws are very different from other analyzed species, thus all observed individuals of this group were found isolated from the others, considering the horizontal axis. Relative warps and curvature analyses show to be useful tools in the forensic study of shark incidents. In general, jaws shape significantly differed among species.

Lost before found: A new species of whaler shark Carcharhinus obsolerus from the Western Central Pacific known only from historic records.

Carcharhinus obsolerus is described based on three specimens from Borneo, Thailand and Vietnam in the Western Central Pacific. It belongs to the porosus subgroup which is characterised by having the second dorsal-fin insertion opposite the anal-fin midbase. It most closely resembles C. borneensis but differs in tooth morphology and counts and a number of morphological characters, including lack of enlarged hyomandibular pores which are diagnostic of C. borneensis. The historic range of C. obsolerus sp. nov. is under intense fishing pressure and this species has not been recorded anywhere in over 80 years. There is an urgent need to assess its extinction risk status for the IUCN Red List of Threatened Species. With so few known records, there is a possibility that Carcharhinus obsolerus sp. nov. has been lost from the marine environment before any understanding could be gained of its full historic distribution, biology, ecosystem role, and importance in local fisheries.

Comparative morphology and systematics of the cookiecutter sharks, genus Isistius Gill (1864) (Chondrichthyes: Squaliformes: Dalatiidae).

The dalatiid genus Isistius Gill (1864) has three valid species currently recognized in the literature: Isistius brasiliensis Quoy & Gaimard (1824), I. plutodus Garrick & Springer (1964), and I. labialis Meng, Zhu & Li (1985). The most common species, I. brasiliensis, has a wide geographic distribution and is found in subtemperate and tropical seas circumglobally. A comparative analysis of specimens from different localities throughout its range, however, had never been undertaken. In the present paper, the morphological variation of this species along its entire distribution has been thoroughly analyzed, corroborating that it represents a single widespread species and that I. labialis is its junior synonym. The other congeneric species, I. plutodus, is known from only a few specimens and is also distributed worldwide. A detailed comparative analysis of available material of I. plutodus was conducted verifying its validity as a single widespread species. The present study analyzed in detail the external morphology (coloration, dentition, dermal denticles), internal morphology (skeleton, musculature), lateral-line canals, and morphometric and meristic characters of species of Isistius in order to better define the genus and its included valid species.

A new microendemic species of the deep-water catshark genus Bythaelurus (Carcharhiniformes, Pentanchidae) from the northwestern Indian Ocean, with investigations of its feeding ecology, generic review and identification key.

A new deep-water catshark, Bythaelurus stewarti, is described based on 121 examined specimens caught on the Error Seamount (Mount Error Guyot) in the northwestern Indian Ocean. The new species differs from all congeners in the restricted distribution, a higher spiral valve turn count and in the morphology of the dermal denticles. It is distinguished from its morphologically and geographically closest congener, B. hispidus (Alcock), by the larger size (maximum size 44 vs. 39 cm TL, maturity size of males 35-39 vs. 21-28 cm TL), darker fresh coloration and dark grayish-brown mottling of the ventral head (vs. ventral head typically uniformly yellowish or whitish). Furthermore, it has a strongly different morphology of dermal denticles, in particular smaller and less elongate branchial, trunk and lateral caudal denticles that are set much less densely and have a surface that is very strongly and fully structured by reticulations (vs. structured by reticulations only in basal fourth). In addition, the new species differs from B. hispidus in having more slender claspers that are gradually narrowing to the bluntly pointed tip without knob-like apex (vs. claspers broader and with distinct knob-like apex), more spiral valve turns (11-12 vs. 8-10) and numerous statistical differences in morphometrics. A review of and a key to the species of Bythaelurus are given.