Tooth development and histology patterns in lamniform sharks (Elasmobranchii,Lamniformes) revisited

The dentition of lamniforme sharks exhibits several characters that have been used extensively to resolve the phylogenetic relationships of extant taxa, yet some uncertainties remain. Also, the development of different teeth of a tooth file within the jaws of most extant lamniforms has not been documented to date. High‐resolution micro‐computed tomography is used here to re‐evaluate the importance of two dental characters within the order Lamniformes, which were considered not to be phylogenetically informative, the histotype and the number of teeth per tooth file. Additionally, the development and mineralization patterns of the teeth of the two osteodont lamniforms Lamna nasus and Alopias superciliosus were compared. We discuss the importance of these dental characters for phylogenetic interpretations to assess the quality of these characters in resolving lamniform relationships. The dental characters suggest that (1) Lamniformes are the only modern‐level sharks exhibiting the osteodont histotype, (2) the osteodont histotype in lamniform sharks is a derived state in modern‐level sharks (Elasmobranchii), (3) the osteodont type, conversely is convergently achieved when the clade Chondrichthyes is considered and thus might comprise a functional rather than a phylogenetic signal, and (4) there is an increase in the number of teeth per file throughout lamniform phylogeny. Structural development of the teeth of L. nasus and A. superciliosus is congruent with a previous investigation of the lamniform shark Carcharodon carcharias. J. Morphol. 277:1584–1598, 2016. © 2016 Wiley Periodicals, Inc.     

A Gigantic Shark from the Lower Cretaceous Duck Creek Formation of Texas

Three large lamniform shark vertebrae are described from the Lower Cretaceous of Texas. We interpret these fossils as belonging to a single individual with a calculated total body length of 6.3 m. This large individual compares favorably to another shark specimen from the roughly contemporaneous Kiowa Shale of Kansas. Neither specimen was recovered with associated teeth, making confident identification of the species impossible. However, both formations share a similar shark fauna, with Leptostyrax macrorhiza being the largest of the common lamniform sharks. Regardless of its actual identification, this new specimen provides further evidence that large-bodied lamniform sharks had evolved prior to the Late Cretaceous.     

Pathologic tooth deformities in modern and fossil chondrichthians: a consequence of feeding-related injury

Deformed teeth are found as rare components of the dentitions of both modern and fossil chondrichthians. Tooth deformities occur as bent or twisted tooth crowns, missing or misshaped cusps, atypical protuberances, perforations, and abnormal root structures. Deformed tooth files consisting of unusually overlapped or small teeth, or teeth misaligned in the jaw also occur in modern forms, but deformed tooth files generally are not recognizable in fossils due to post-mortem dissociation of teeth and jaws. A survey of 200 modern lamniform and carcharhiniform sharks as well as literature sources indicate that such deformities are produced by feeding-related injury to the tooth-forming tissue of the jaws, particularly by impaction of chondrichthian and teleost fin and tail spines. Tooth counts for several late Cretaceous genera, based on material recovered from coastal plain sites from New Jersey to Alabama, suggest that the frequency of occurrence of deformed teeth in a species varies from about 0.015% in Squalicorax kaupi to about 0.36% in Paranomotodon sp. Tooth counts for modern lamniform and carcharhiniform sharks yield a comparable range in frequency of tooth deformities. Variation in frequency of tooth deformity may reflect interspecific differences in feeding behavior and dietary preferences. There is no suggestion in our data of any strong patterns of temporal variation in tooth deformity frequency, or of patterns ­reflecting chondrichthian phylogenetic history and evolution. Skeletal components of the probable prey of the Cretaceous species are preserved in the same horizons as the deformed teeth, and also are found within co-occurring chondrichthian coprolites.     

A new Early Cretaceous lamniform shark (Chondrichthyes,Neoselachii)

Eoptolamna eccentrolopha gen. et sp. nov. (Chondrichthyes, Lamniformes) from the near coastal upper Barremian Artoles Formation (Early Cretaceous) of Castellote (northwestern Spain) is described on the basis of about 50 isolated teeth. This taxon represents one of the earliest lamniform sharks known to date. We hypothesize that most pre‐Aptian lamniforms belong to an ancient group characterized, amongst others, by a very weak gradient monognathic heterodont dental pattern, and by tearing‐type dentition. There is a nutritive groove in the lingual root protuberance in juveniles of Eoptolamna, which persists in adults. A single pair of symphysial and a pair of upper intermediate teeth might have been present. Consequently, a new family, Eoptolamnidae, is introduced to include the new form, as well as Protolamna and probably Leptostyrax. The Eoptolamnidae represent an ancient family within Lamniformes. The origin of lamniform sharks remains, however, ambiguous despite recent advances. The new Spanish taxon is widespread in the Barremian of north‐eastern Spain, and occurs in a wide range of facies from near‐coastal to lake deposits. This lamniform also occurs in the Lower Cretaceous of northern Africa. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 278–290.     

Late Cretaceous sharks from Cuba,first record of Serratolamna serrata (Agassiz) (Lamniformes,Serratolamnidae)

Only one species of Elasmobranchii, Ptychodus cyclodontis Mutter, Iturralde-Vinent and Carmona (2005), has been reported so far from the Late Cretaceous of Cuba. Herein we describe the first record of a Maastrichtian Serratolamna serrata (Agassiz, 1843) as well as non-diagnostic remains which include a tooth referred to a lamniform shark and an isolated vertebra of an indeterminate elasmobranch. These fossils expand the temporal distribution of Cretaceous fossil sharks known from Cuba and increase our understanding of the group’s fossil diversity.     

Dental ontogeny of a white shark embryo

Unlike most viviparous vertebrates, lamniform sharks develop functional teeth during early gestation. This feature is considered to be related to their unique reproductive mode where the embryo grows to a large size via feeding on nutritive eggs in utero. However, the developmental process of embryonic teeth is largely uninvestigated. We conducted X‐ray microcomputed tomography to observe the dentitions of early‐, mid‐, and full‐term embryos of the white shark Carcharodon carcharias (Lamniformes, Lamnidae). These data reveal the ontogenetic change of embryonic dentition of the species for the first time. Dentition of the early‐term embryos (∼45 cm precaudal length, PCL) is distinguished from adult dentition by 1) the presence of microscopic teeth in the distalmost region of the paratoquadrate, 2) a fang‐like crown morphology, and 3) a lack of basal concavity of the tooth root. The “intermediate tooth” of early‐term embryos is almost the same size as the adjacent teeth, suggesting that lamnoid‐type heterodonty (lamnoid tooth pattern) has not yet been established. We also discovered that mid‐term embryos (∼80 cm PCL) lack functional dentition. Previous studies have shown that the maternal supply of nutritive eggs in lamnoid sharks ceases during mid‐ to late‐gestation. Thus, discontinuation of functional tooth development is likely associated with the completion of the oophagous (egg‐eating) phase. Replacement teeth in mid‐term embryos include both embryonic and adult‐type teeth, suggesting that the embryo to adult transition in dental morphology occurs during this period. J. Morphol. 278:215–227, 2017. © 2016 Wiley Periodicals,Inc.     

Teeth of Embryos in Lamniform Sharks (Chondrichthyes: Elasmobranchii)

The dentitions of lamniform sharks possess a unique heterodonty, the lamnoid tooth pattern. However, in embryos, there are 'embryonic' and 'adult' dentitions. The teeth in the embryonic dentition are peg-like and appear to be attached to the jaw in an acrodont fashion. The adult dentition is characterized by the presence of replacement tooth series with the lamnoid tooth pattern. The embryonic–adult transition in dentitions appears at around 30–60cm TL. Tooth replacement generally begins before birth in embryos with adult dentitions. The adult dentition becomes functional just before or after parturition. An embryo of one species (Lamna nasus) shows a tooth directly on the symphysis of the upper jaws, marking the first record of a medial tooth for the order Lamniformes.     

A new assemblage of freshwater sharks (Chondrichthyes: Elasmobranchii) from the Upper Triassic of India

This study reports the first occurrence of a varied xenacanth assemblage from the Upper Triassic Tiki Formation of India, based on multiple well-preserved isolated teeth. Based on distinct tooth morphology, two species of the genus Mooreodontus are described: M. indicus and a new species, M. jaini. The new species is diagnosed based on a tricuspid crown containing two stout, slightly diverging lateral cusps pointing in the same direction, a high median cusp, crown-base angle almost at 90°, large, rounded, apical button with several foramina and multiple, 8–9 coarse vertical cristae on all the cusps. Dental anomaly in the form of a partial quadri-cuspidate xenacanthid tooth is present in the collection. Another group of xenacanthid teeth have bicuspid crowns with two upright, asymmetric cusps, where the mesial cusp is thicker than the distal one, and consistently lack a median cusp. Such distinct bicuspid tooth morphology is usually present in Palaeozoic forms and is reported for the first time from the Late Triassic. It is considered to belong to a new taxon, Tikiodontus asymmetricus nov. gen., nov. sp., of indeterminate family. Distinctive tooth histology also differentiates the two Indian genera Mooreodontus and Tikiodontus nov. gen. from other xenacanthid taxa. In addition, the Tiki assemblage has yielded multiple chondrichthyan dermal denticles, which may be subdivided into two morphotypes based on their robustness and presence/absence of linear ridges on the fused cusps. India holds a unique position in terms of its Late Triassic freshwater shark fauna, as it exhibits distinct Laurasian affinities. These freshwater sharks had restricted occurrences in other parts of the Gondwanan landmass.     

The Upper Jurassic selachian Palaeocarcharias de Beaumont (1960)

The teeth of Palaeocarcharias stromeri De Beaumont (Kimmeridgian, Upper Jurassic; southern Germany) show linear gradient monognathic heterodonty from high-cusped parasymphyseal teeth to low-crowned posterolaterals. The bifid root has holaulacorhize vascularization and is surmounted by mesial and distal extensions of the triple-layered coronal enameloid. The crowns lack lateral cusplets, but possess short vertical striations at the base of the central cusp labially and lingually. A collar is developed at the base of the central cusp lingually, and a small node in a similar position labially. It is concluded that Palaeocarcharias must represent the oldest known lamniform. The body form of Palaeocarcharias is most closely comparable to that of the orectolobiform sharks, but the teeth are lamnoid. Palaeocarcharias is best interpreted as a benthonic stem group lamniform. The lamniforms are visualized as arising from orectolobiform ancestors, probably in the middle Jurassic.     

Dental homologies in lamniform sharks (Chondrichthyes: Elasmobranchii).

The dentitions of lamniform sharks are said to exhibit a unique heterodonty called the "lamnoid tooth pattern." The presence of an inflated hollow "dental bulla" on each jaw cartilage allows the recognition of homologous teeth across most modern macrophagous lamniforms based on topographic correspondence through the "similarity test." In most macrophagous lamniforms, three tooth rows are supported by the upper dental bulla: two rows of large anterior teeth followed by a row of small intermediate teeth. The lower tooth row occluding between the two rows of upper anterior teeth is the first lower anterior tooth row. Like the first and second lower anterior tooth rows, the third lower tooth row is supported by the dental bulla and may be called the first lower intermediate tooth row. The lower intermediate tooth row occludes between the first and second upper lateral tooth rows situated distal to the upper dental bulla, and the rest of the upper and lower tooth rows, all called lateral tooth rows, occlude alternately. Tooth symmetry cannot be used to identify their dental homology. The presence of dental bullae can be regarded as a synapomorphy of Lamniformes and this character is more definable than the "lamnoid tooth pattern." The formation of the tooth pattern appears to be related to the evolution of dental bullae. This study constitutes the first demonstration of supraspecific tooth-to-tooth dental homologies in nonmammalian vertebrates.     

Teeth of enigmatic neoselachian sharks and an ornithischian dinosaur from the uppermost Triassic of Lons-le-Saunier (Jura, France)

Shark teeth and an ornithischian dinosaur tooth are described from a new palynologically dated Rhaetian locality at Lons-le-Saunier (Jura, France). The structure of the enameloid of the teeth ofSynechodus rhaeticus has been studied, but this appears quite different from the usual pattern seen in neoselachian sharks, making the precise relationships of this species difficult to determine. On the other hand,‘Hybodus’ minor, which has long be thought to be a hybodont shark, is included among the Synechodontiformes. The find of the tooth of an ornithischian dinosaur is also reported. Study of the Lonsle-Saunier site seems to indicate a change in the marine faunas during the Rhaetian transgression, preferentially affecting the neoselachian sharks, which increase in abundance, and thedurophasous bony fishes, which become dominated bySareodon tomicus.     

Theropod dinosaur teeth from the lowermost Cretaceous Rabekke Formation on Bornholm, Denmark

The dinosaur fauna of the palynologically dated lower Berriasian Skyttegård Member of the Rabekke Formation on the Baltic island of Bornholm, Denmark, is represented by isolated tooth crowns. The assemblage is restricted to small maniraptoran theropods, assigned to the Dromaeosauridae incertae sedis and Maniraptora incertae sedis. The dromaeosaurid teeth are characterized by their labiolingually compressed and distally curved crowns that are each equipped with a lingually flexed mesial carina and a distinctly denticulated distal cutting edge. A morphologically aberrant tooth crown (referred to as Maniraptora incertae sedis) has triangular denticles of uneven width, a feature occasionally found in Upper Cretaceous hesperornithiform toothed diving birds, but also in premaxillary teeth of the velociraptorine Nuthetes from the Lower Cretaceous of England.     

Systematics,palaeoecology and palaeobiogeography of the Neogene fossil sharks from the Azores (Northeast Atlantic)

The Azores Archipelago is a group of isolated islands located in the North Atlantic Ocean. One of these oceanic islands – Santa Maria – exhibits marine fossiliferous sediments of late Miocene/early Pliocene and also of Pleistocene age. Recent research provided new selachian fossil material, with three new records (Carcharias acutissima, Megaselachus megalodon, and Carcharhinus cf. leucas) increasing the number of fossil sharks reported from the Azores (Santa Maria Island) to seven species (Notorynchus primigenius, C. acutissima, Cosmopolitodus hastalis, Paratodus benedenii, Isurus oxyrinchus, M. megalodon, and C. cf. leucas). So far, no teeth of batoids or small sharks have been found despite the screen-washing of several sediment samples from Santa Maria. The Azorean Mio-Pliocene selachian fauna clearly differs from those described from sediments deposited on continental shelves, in which batoids and small benthic sharks (e.g., scyliorhinids) are usually well represented. During the late Miocene/early Pliocene, subtropical to warm-temperate seas were prevalent in the area of the Azores, as deduced from palaeontological, geological and isotopic data, all indicating a warmer climate than in the present.     

Contributions to the tooth morphology in early embryos of three species of hammerhead sharks (Elasmobranchii: Sphyrnidae) and their evolutionary implications

The tooth types in the embryos of the hammerhead sharks Sphyrna tiburo, Sphyrna tudes and Eusphyra blochii are here described in labial and lingual views, and, in some cases, in additional views. The presence of cusplets was observed in the anterior teeth of S. tiburo and S. tudes, which is secondarily lost after early embryonic stages. Many aligned root foramina were detected in the sphyrnids, which, as the cusplets, are shared by many phylogenetic-related carcharhinids. Other anatomic features, related to the root and central cusp, are presented for the first time. Such characters represent the first step to compare the teeth of extant and fossil species.     

Basilotritus (Cetacea: Pelagiceti) from the Eocene of Nagornoye (Ukraine): New data on anatomy,ontogeny and feeding of early basilosaurids

A new specimen of early basilosaurid, now identified as Basilotritus sp., comes from the late Middle Eocene of Ukraine. It has basilosaurid-type cheek teeth with cinguli, similar to those of Zygorhiza, and roots resembling those of Georgiacetus vogtlensis and early Neoceti; an unusual feature of these teeth is the presence of accessory denticles of the second order located on the crown denticles. The postcranial anatomy shows a mixture of primitive and advanced basilosaurid traits. The phylogenetic position of the genus Basilotritus is confirmed to be near the base of Basilosauridae, between Supayacetus and Zygorhiza. The ontogeny of the specimen from Nagornoye is characterized by large body size, slow skeletal maturation and intensive pachyosteosclerosis that are interpreted as neotenic development. Rapid tooth wear with strong apical abrasion is the result of specialized diet, possibly feeding on sharks.     

Tooth morphology elucidates shark evolution across the end-Cretaceous mass extinction

Sharks (Selachimorpha) are iconic marine predators that have survived multiple mass extinctions over geologic time. Their prolific fossil record is represented mainly by isolated shed teeth, which provide the basis for reconstructing deep time diversity changes affecting different selachimorph clades. By contrast, corresponding shifts in shark ecology, as measured through morphological disparity, have received comparatively limited analytical attention. Here, we use a geometric morphometric approach to comprehensively examine tooth morphologies in multiple shark lineages traversing the catastrophic end-Cretaceous mass extinction—this event terminated the Mesozoic Era 66 million years ago. Our results show that selachimorphs maintained virtually static levels of dental disparity in most of their constituent clades across the Cretaceous–Paleogene interval. Nevertheless, selective extinctions did impact apex predator species characterized by triangular blade-like teeth. This is particularly evident among lamniforms, which included the dominant Cretaceous anacoracids. Conversely, other groups, such as carcharhiniforms and orectolobiforms, experienced disparity modifications, while heterodontiforms, hexanchiforms, squaliforms, squatiniforms, and †synechodontiforms were not overtly affected. Finally, while some lamniform lineages disappeared, others underwent postextinction disparity increases, especially odontaspidids, which are typified by narrow-cusped teeth adapted for feeding on fishes. Notably, this increase coincides with the early Paleogene radiation of teleosts as a possible prey source, and the geographic relocation of disparity sampling “hotspots,” perhaps indicating a regionally disjunct extinction recovery. Ultimately, our study reveals a complex morphological response to the end-Cretaceous mass extinction and highlights an event that influenced the evolution of modern sharks.

Analysis of the tooth morphology of sharks across the end-Cretaceous mass extinction, 66 million years ago, shows that while generally unaffected, some apex predator shark lineages were selectively impacted; changing habitats and the differential survival of ‘fish-eating’ sharks also reveals responses to ecological cataclysm.     

Palaeoenvironments of Early Miocene hominoid-bearing deposits at Napak,Uganda, based on terrestrial molluscs

Studies of fossil molluscs from Early Miocene deposits at Napak, Uganda, reveal that between 20 and 18.5 Ma, there were various kinds of palaeoenvironments on the flanks of the volcano ranging from forest to open country. The basal Iriri Member contains fluvial deposits from which freshwater gastropods and bivalves have been collected. The younger Napak Member is subaerial in facies, and has yielded a predominantly forest assemblage of gastropods, but with some indications of open country species at one of the sites. Napak has yielded a rich and diverse primate fauna, including the large ape Ugandapithecus major, the small ape Micropithecus clarki, the earliest known cercopithecid in the world (Prohylobates sp.) and several galagids (Mioeuoticus bishopi) among other less well-known taxa.     

Pathologic tooth deformities in fossil and modern sharks related to jaw injuries

Dental abnormalities in a tiger shark Galeocerdo cuvier and in Carcharoides totuserratus are presented here again, along with some further ones in shark teeth. Comparisons are made with fossil and modern shark teeth abnormalities. A coalescent set of two Squalicorax pristodontus teeth is described. The gibbous shape of the crown is similar to that in S. kaupi, the preceding species of the same lineage. It therefore suggests that the differentiation of the most derived species S. pristodontus may have resulted from kaupi through size increase and development of jaws, becoming more spacious, with teeth getting broader mesio-distally. An abnormal Carcharocles megalodon tooth is described. We regard it as a left lateral tooth from the mandible, whose crown is much deformed. Its features suggest trauma resulting from a feeding accident, maybe through biting the very compact bones of its more likely common prey, Halianassa sirenians. The last case concerns an abnormal Negaprion tooth. The most remarkable differences apart from the normal teeth concern the crown, which is irregular in shape. It shows some torsion, which also occurs in the root. A well-marked notch occurs in the mesial side. The cusp is somewhat labially bent. Trauma on the tooth-forming tissues seems to be responsible for the abnormalities under study. As far as we could ascertain, no lemon-shark dental abnormalities have previously been described. Our results stress that tooth modifications resulting from injuries to the tooth-producing tissues occurred since long ago in similar ways as in extant sharks. Biting prey's hard skeletal parts seems as always the main cause for injuries.     

New fossils of Australopithecus anamensis from Kanapoi,West Turkana,Kenya (2003–2008)

Renewed fieldwork from 2003 through 2008 at the Australopithecus anamensis type-site of Kanapoi, Kenya, yielded nine new fossils attributable to this species. These fossils all date to between 4.195 and 4.108 million years ago. Most were recovered from the lower fluvial sequence at the site, with one from the lacustrine sequence deltaic sands that overlie the lower fluvial deposits but are still below the Kanapoi Tuff. The new specimens include a partial edentulous mandible, partial maxillary dentition, two partial mandibular dentitions, and five isolated teeth. The new Kanapoi hominin fossils increase the sample known from the earliest Australopithecus, and provide new insights into morphology within this taxon. They support the distinctiveness of the early A. anamensis fossils relative to earlier hominins and to the later Australopithecus afarensis. The new fossils do not appreciably extend the range of observed variation in A. anamensis from Kanapoi, with the exception of some slightly larger molars, and a canine tooth root that is the largest in the hominin fossil record. All of the Kanapoi hominins share a distinctive morphology of the canine–premolar complex, typical early hominin low canine crowns but with mesiodistally longer honing teeth than seen in A. afarensis, and large, probably dimorphic, canine tooth roots. The new Kanapoi specimens support the observation that canine crown height, morphology, root size and dimorphism were not altered from a primitive ape-like condition as part of a single event in human evolution, and that there may have been an adaptive difference in canine function between A. anamensis and A. afarensis.