Enameloid Research Articles

The late Cenozoic evolution of the Humboldt Current System in coastal Peru: Insights from neodymium isotopes

The Humboldt Current System along the Pacific coast of South America creates one of the most productive ecosystems on Earth. To trace the origin of the water masses in this area, we measured neodymium isotope compositions (ԑNd) in tooth enameloid of two genera of coastal sharks from latest Oligocene to early Pleistocene strata in the Pisco and Sacaco basins in southern Peru. Most ԑNd values range from −4 to −1, with a strong negative excursion in the late Miocene (∼8–7 million years ago [Ma]) with values as low as −9.2. The overall trend of the ԑNd values resembles that of equatorial Pacific deep waters, though with an offset of about +2 ԑNd units until about 6 Ma. With a major input of hinterland weathering considered unlikely, we interpret this pattern as reflecting a modern-type upwelling regime, though with a lower contribution of Antarctic waters than today. Starting about 6 Ma, the contribution of Antarctic waters to the upwelling waters increased approximately to present-day levels, coincident with, and possibly driven by, increased Antarctic glaciation and the Andes reaching their present-day elevation, both of which likely enhanced the counter-clockwise circulation in the South Pacific Ocean. The negative excursion of ԑNd values in the Pisco/Sacaco basins ∼8–7 Ma coincides with a late Miocene biogenic bloom in the Pacific Ocean and elsewhere, and with a strongly increased northward bottom current observed on the Nazca Drift System just offshore our sampling area. Thus, the negative excursion of ԑNd values in the Pisco/Sacaco basins likely resulted from a southern sourced input of nutrient-rich, unradiogenic water, which could have been an important contributor to the biogenic bloom.

Trophic position of Otodus megalodon and great white sharks through time revealed by zinc isotopes

Diet is a crucial trait of an animal’s lifestyle and ecology. The trophic level of an organism indicates its functional position within an ecosystem and holds significance for its ecology and evolution. Here, we demonstrate the use of zinc isotopes (δ66Zn) to geochemically assess the trophic level in diverse extant and extinct sharks, including the Neogene megatooth shark (Otodus megalodon) and the great white shark (Carcharodon carcharias). We reveal that dietary δ66Zn signatures are preserved in fossil shark tooth enameloid over deep geologic time and are robust recorders of each species’ trophic level. We observe significant δ66Zn differences among the Otodus and Carcharodon populations implying dietary shifts throughout the Neogene in both genera. Notably, Early Pliocene sympatric C. carcharias and O. megalodon appear to have occupied a similar mean trophic level, a finding that may hold clues to the extinction of the gigantic Neogene megatooth shark.

Hierarchical Microstructure of Tooth Enameloid in Two Lamniform Shark Species, Carcharias taurus and Isurus oxyrinchus.

Shark tooth enameloid is a hard tissue made up of nanoscale fluorapatite crystallites arranged in a unique hierarchical pattern. This microstructural design results in a macroscopic material that is stiff, strong, and tough, despite consisting almost completely of brittle mineral. In this contribution, we characterize and compare the enameloid microstructure of two modern lamniform sharks, Isurus oxyrinchus (shortfin mako shark) and Carcharias taurus (spotted ragged-tooth shark), based on scanning electron microscopy images. The hierarchical microstructure of shark enameloid is discussed in comparison with amniote enamel. Striking similarities in the microstructures of the two hard tissues are found. Identical structural motifs have developed on different levels of the hierarchy in response to similar biomechanical requirements in enameloid and enamel. Analyzing these structural patterns allows the identification of general microstructural design principles and their biomechanical function, thus paving the way for the design of bioinspired composite materials with superior properties such as high strength combined with high fracture resistance.

Fossil microbial shark tooth decay documents in situ metabolism of enameloid proteins as nutrition source in deep water environments

Alteration of organic remains during the transition from the bio- to lithosphere is affected strongly by biotic processes of microbes influencing the potential of dead matter to become fossilized or vanish ultimately. If fossilized, bones, cartilage, and tooth dentine often display traces of bioerosion caused by destructive microbes. The causal agents, however, usually remain ambiguous. Here we present a new type of tissue alteration in fossil deep-sea shark teeth with in situ preservation of the responsible organisms embedded in a delicate filmy substance identified as extrapolymeric matter. The invading microorganisms are arranged in nest- or chain-like patterns between fluorapatite bundles of the superficial enameloid. Chemical analysis of the bacteriomorph structures indicates replacement by a phyllosilicate, which enabled in situ preservation. Our results imply that bacteria invaded the hypermineralized tissue for harvesting intra-crystalline bound organic matter, which provided nutrient supply in a nutrient depleted deep-marine environment they inhabited. We document here for the first time in situ bacteria preservation in tooth enameloid, one of the hardest mineralized tissues developed by animals. This unambiguously verifies that microbes also colonize highly mineralized dental capping tissues with only minor organic content when nutrients are scarce as in deep-marine environments.

Probing the Ecology and Climate of the Eocene Southern Ocean With Sand Tiger Sharks Striatolamia macrota.

Many explanations for Eocene climate change focus on the Southern Ocean—where tectonics influenced oceanic gateways, ocean circulation reduced heat transport, and greenhouse gas declines prompted glaciation. To date, few studies focus on marine vertebrates at high latitudes to discern paleoecological and paleoenvironmental impacts of this climate transition. The Tertiary Eocene La Meseta (TELM) Formation has a rich fossil assemblage to characterize these impacts; Striatolamia macrota, an extinct (†) sand tiger shark, is abundant throughout the La Meseta Formation. Body size is often tracked to characterize and integrate across multiple ecological dimensions. †S. macrota body size distributions indicate limited changes during TELMs 2–5 based on anterior tooth crown height (n = 450, mean = 19.6 ± 6.4 mm). Similarly, environmental conditions remained stable through this period based on δ18OPO4 values from tooth enameloid (n = 42; 21.5 ± 1.6‰), which corresponds to a mean temperature of 22.0 ± 4.0°C. Our preliminary ε Nd (n = 4) results indicate an early Drake Passage opening with Pacific inputs during TELM 2–3 (45–43 Ma) based on single unit variation with an overall radiogenic trend. Two possible hypotheses to explain these observations are (1) †S. macrota modified its migration behavior to ameliorate environmental changes related to the Drake Passage opening, or (2) the local climate change was small and gateway opening had little impact. While we cannot rule out an ecological explanation, a comparison with climate model results suggests that increased CO2 produces warm conditions that also parsimoniously explain the observations.

Shape-preserving erosion controlled by the graded microarchitecture of shark tooth enameloid

The teeth of all vertebrates predominantly comprise the same materials, but their lifespans vary widely: in stark contrast to mammals, shark teeth are functional only for weeks, rather than decades, making lifelong durability largely irrelevant. However, their diets are diverse and often mechanically demanding, and as such, their teeth should maintain a functional morphology, even in the face of extremely high and potentially damaging contact stresses. Here, we reconcile the dilemma between the need for an operative tooth geometry and the unavoidable damage inherent to feeding on hard foods, demonstrating that the tooth cusps of Port Jackson sharks, hard-shelled prey specialists, possess unusual microarchitecture that controls tooth erosion in a way that maintains functional cusp shape. The graded architecture in the enameloid provokes a location-specific damage response, combining chipping of outer enameloid and smooth wear of inner enameloid to preserve an efficient shape for grasping hard prey. Our discovery provides experimental support for the dominant theory that multi-layered tooth enameloid facilitated evolutionary diversification of shark ecologies.

A Lower Cretaceous chondrichthyan dermal denticle assemblage and its bearing on placoid scale diversity and histology

Dermal denticles of Mesozoic chondrichthyans are rarely documented, which tremendously limits the knowledge of different morphotypes during this time and for the whole clade. Here we describe an extraordinarily diverse assemblage of dermal denticles from deposits of Austria, comprising 17 different morphotypes. Based on the functional types of the denticles and their morphological similarities with both extinct and extant representatives, we attempt an order-level taxonomic attribution of the morphotypes recovered. The herein recorded occurrences of Hexanchiformes, Squaliformes, Squatiniformes, Heterodontiformes, the family Protospinacidae (Squalomorphii incertae sedis), and probably Rajiformes or Chimaeriformes increase previous tooth-based taxic diversity estimates of Valanginian cartilaginous fishes from Austria from four to at least nine orders. Additionally, we discuss the absence of denticles of cladodontomorph chondrichthyans, whose teeth were previously reported from the same deposits. Etched sections and light microscope analysis were carried out using the most abundant morphotypes with the aim of exploring the histology of chondrichthyan dermal denticles. Our results confirm previous studies in reporting a single crystallite enameloid, but we also report for the first time the presence of bundled enameloid in dermal scales, a condition so far restricted to the tooth enameloid of some chondrichthyans. Additionally, a putative new dentine type is observed, leading to the introduction of the descriptive term ‘nepedentine’ for the internal, dense, and poorly supplied dentine core by vascular channels.

Past aquatic environments in the Levant inferred from stable isotope compositions of carbonate and phosphate in fish teeth.

Here we explore the carbon and oxygen isotope compositions of the co-existing carbonate and phosphate fractions of fish tooth enameloid as a tool to reconstruct past aquatic fish environments and harvesting grounds. The enameloid oxygen isotope compositions of the phosphate fraction (δ18OPO4) vary by as much as ~4‰ for migratory marine fish such as gilthead seabream (Sparus aurata), predominantly reflecting the different saline habitats it occupies during its life cycle. The offset in enameloid Δ18OCO3-PO4 values of modern marine Sparidae and freshwater Cyprinidae from the Southeast Mediterranean region vary between 8.1 and 11.0‰, similar to values reported for modern sharks. The mean δ13C of modern adult S. aurata and Cyprinus carpio teeth of 0.1±0.4‰ and -6.1±0.7‰, respectively, mainly reflect the difference in δ13C of dissolved inorganic carbon (DIC) of the ambient water and dietary carbon sources. The enameloid Δ18OCO3-PO4 and δ13C values of ancient S. aurata (Holocene) and fossil Luciobarbus sp. (Cyprinidae; mid Pleistocene) teeth agree well with those of modern specimens, implying little diagenetic alteration of these tooth samples. Paired δ18OPO4-δ13C data from ancient S. aurata teeth indicate that hypersaline water bodies formed in the Levant region during the Late Holocene from typical Mediterranean coastal water with high evaporation rates and limited carbon input from terrestrial sources. Sparid tooth stable isotopes further suggest that coastal lagoons in the Eastern Mediterranean had already formed by the Early Holocene and were influenced by terrestrial carbon sources. Overall, combined enameloid oxygen and carbon isotope analysis of fish teeth is a powerful tool to infer the hydrologic evolution of aquatic environments and assess past fishing grounds of human populations in antiquity.

Tooth oxygen isotopes reveal Late Bronze Age origin of Mediterranean fish aquaculture and trade

Past fish provenance, exploitation and trade patterns were studied by analyzing phosphate oxygen isotope compositions (δ18OPO4) of gilthead seabream (Sparus aurata) tooth enameloid from archaeological sites across the southern Levant, spanning the entire Holocene. We report the earliest evidence for extensive fish exploitation from the hypersaline Bardawil lagoon on Egypt’s northern Sinai coast, as indicated by distinctively high δ18OPO4 values, which became abundant in the southern Levant, both along the coast and further inland, at least from the Late Bronze Age (3,550–3,200 BP). A period of global, postglacial sea-level stabilization triggered the formation of the Bardawil lagoon, which was intensively exploited and supported a widespread fish trade. This represents the earliest roots of marine proto-aquaculture in Late Holocene coastal domains of the Mediterranean. We demonstrate the potential of large-scale δ18OPO4 analysis of fish teeth to reveal cultural phenomena in antiquity, providing unprecedented insights into past trade patterns.

Immunolocalization of enamel matrix protein-like proteins in the tooth enameloid of spotted gar, Lepisosteus oculatus, an actinopterygian bony fish

ABSTRACTEnameloid is a well-mineralized tissue covering the tooth surface in fish and it corresponds to the outer-most layer of dentin. It was reported that both dental epithelial cells and odontoblasts are involved in the formation of enameloid. Nevertheless, the localization and timing of secretion of ectodermal enamel matrix proteins in enameloid are unclear. In the present study, the enameloid matrix during the stages of enameloid formation in spotted gar, Lepisosteus oculatus, an actinopterygian, was examined mainly by transmission electron microscopy-based immunohistochemistry using an anti-mammalian amelogenin antibody and antiserum. Positive immunoreactivity with the antibody and antiserum was found in enameloid from the surface to the dentin-enameloid junction just before the formation of crystallites. This immunoreactivity disappeared rapidly before the full appearance of crystallites in the enameloid during the stage of mineralization. Immunolabelling was usually found along the collagen fibrils but was not seen on the electron-dense fibrous structures, which were probably derived from matrix vesicles in the previous stage. In inner dental epithelial cells, the granules in the distal cytoplasm often showed positive immunoreactivity, suggesting that the enamel matrix protein-like proteins originated from inner dental epithelial cells. Enamel matrix protein-like proteins in the enameloid matrix might be common to the enamel matrix protein-like proteins previously reported in the collar enamel of teeth and ganoine of ganoid scales, because they exhibited marked immunoreactivity with the same anti-mammalian amelogenin antibodies. It is likely that enamel matrix protein-like proteins are involved in the formation of crystallites along collagen fibrils in enameloid.

Fluor‑carbonated hydroxyapatite coatings by pulsed laser deposition to promote cell viability and antibacterial properties

Fluorine, an essential element present in bone and dental tissues, promotes mineralization and is directly involved in the bone formation process. The antibacterial effects of fluorine on oral bacteria are also well known. In this study, metallic implants were coated with a thin layer of calcium phosphate enriched with fluorine to improve osteointegration and protect against infections. These coatings were obtained by pulsed laser deposition (PLD) using, as a target, a bioceramic of marine origin, mainly composed of fluorapatite from the enameloid of shark teeth. The compositional dependence of coatings on H2O vapor pressures applied during PLD was analyzed in order to optimize physicochemical properties. Physicochemical characterization to evaluate morphology (SEM), thickness (interferometric profilometry), structure (XRD) and composition (FTIR, XPS) was performed. To evaluate the biological response, both MC3T3-E1 pre-osteoblasts and bacterial strains Staphylococcus aureus and Staphylococcus epidermidis, most responsible for 77% of infections associated with prosthetic implants, were tested. Proper cell proliferation (MTT assay) and ALP synthesis up to 21 days were confirmed. Antibacterial properties were also demonstrated: compared to synthetic hydroxyapatite coatings, there was a significant reduction in colony-forming units (CFUs) for both strains.

Glauconite-bearing sedimentary phosphorites from the Tébessa region (eastern Algeria): Evidence of REE enrichment and geochemical constraints on their origin

Rare earth element (REE) analyses are reported on glauconite-bearing phosphorites from northeastern Algeria. These rocks of Paleocene-Eocene age are located in the Eastern part of the Saharan Atlas, where two localities were investigated: the Djebel El Kouif in the north and the Kef Essenoun in the south. The latter belongs to the world-class Djebel Onk mining deposit. Petrographic examination indicates that phosphorite from the main layer of the Kef Essenoun deposit has a significant abundance in glauconite grains, while their occurrence in the Djebel El Kouif is restricted to the basal levels. In both deposits, glauconite grains are richer in REEs than other co-existing particles (pellets, coprolites, enameloid and dentine of marine fish teeth), but the glauconites of the Kef Essenoun deposit exhibit significantly higher REE concentration (min = 654 ppm, max = 1760 ppm, average = 1146 ppm) than those of Djebel El Kouif deposit (min = 543 ppm, max = 623 ppm, average = 584 ppm). The whole-rock REE concentrations also show substantial differences between the two deposits and the REE enrichment in the Kef Essenoun main sub-layer is more likely the result of the high glauconite content. PAAS normalized-REE patterns of the glauconite grains display similar patterns between the northern and southern localities with weak negative Ce anomalies and slight middle REE enrichments. On the other hand, the pellets and whole phosphorite grains from the northern deposit indicate a REE source from oxic-suboxic seawater, whereas those from the Kef Essenoun deposit exhibit slight middle REE enriched patterns with weak negative Ce anomaly suggesting a different environment of deposition (i.e. a tendency to reduced conditions). These geochemical results along with previous petrographic studies confirm the allochthonous character of the main phosphorite sub-layer in Kef Essenoun, where winnowing, transport and re-deposition of previously deposited phosphorites resulted in high accumulation and formation of glauconite grains. The glauconitization process of fecal pellets happened in two stages and was controlled by semi-confined micro-environments. This can explain the abundance of glauconite grains in the phosphorites from Kef Essenoun deposit and their restricted occurrence in the basal levels of the Djebel El Kouif outcrop.

Scanning electron microscope examination of the dental enameloid of the Cretaceous durophagous sharkPtychodussupports neoselachian classification

AbstractAlthoughPtychodusteeth are well known in Late Cretaceous marine deposits in North America and Europe and a few specimens with jaw elements have been discovered, the taxonomic position of the shark genusPtychodusis enigmatic due to the lack of preservation of diagnostic material other than teeth. These sharks possessed a pavement dentition suited to a diet of hard-shelled macroinvertebrates (durophagy), leading several studies to variously describePtychodusas a batoid, a hybodont shark, or a selachimorph. Members of the Selachimorpha share one dental synapomorphy, a triple-layered enameloid (TLE) consisting of an outer shiny-layered enameloid (SLE) of randomly oriented hydroxyapatite crystallites, a middle layer of parallel-bundled enameloid (PBE), and an inner layer of tangled-bundled enameloid (TBE). Batoids and hybodonts both have teeth with single crystallite enameloid (SCE). We examined teeth fromPtychoduscollected from the Lincoln Limestone of the Greenhorn Formation of Barton County, Kansas, and compared their enameloid ultrastructure with that of a Carboniferous hybodontiform and the Cretaceous lamniform sharkSqualicorax curvatusWilliston, 1900. Scanning electron microscopic examination ofPtychodusshows that crystallite bundling in the form of a TLE is evident in these teeth. The PBE is most apparent at transverse enameloid ridges ofPtychodusteeth. Columns of dentine penetrate into the tooth enameloid, and the crystallites near the dentine are randomly oriented. These observations bolster the argument thatPtychodusis a genus of highly specialized selachimorph shark, rather than a hybodont or batoid.

The improved biological response of shark tooth bioapatites in a comparative in vitro study with synthetic and bovine bone grafts

Autologous bone is considered to be the gold standard for bone tissue regeneration, providing more highly efficient functional responses compared to synthetic materials, and avoiding the rejection risks of allogenic grafts. However, it presents limitations for certain types of surgery due to its high resorption levels and donor site morbidity. Different biphasic synthetic composites, based onnon-apatitic calcium phosphates enriched with apatitic phases—such as hydroxyapatite, and bioderived bone grafts of bovine and porcine origin—are proposed as lower resorption materials due to their higher crystalline structure. The present work proposes two new sources of bioapatites for bone filler applications obtained from the dentine and enameloid of shark teeth, respectively. These bioapatites each present a characteristic apatite-based composition and additional enrichments of specific trace elements, such as magnesium and fluorine, with proven roles in bone metabolism. Their processing and physicochemical characterization (SEM, FT-Raman and XRD) is presented, together with an in vitro evaluation of osteogenic activity compared to a commercial bovine mineralized matrix and synthetic HA/β TCP grafts. The results proved the globular morphology (0.5–1.5 μm) and porosity (~50 μm and ~0.5–1 μm) of shark dentine bioapatites with biphasic composition: apatitic (hydroxyapatite and apatite-(CaF)), non-apatitic (whitlockite), and an apatitic phase (fluorapatite), organized in oriented crystals in enameloid bioapatites. An evaluation of the pre-osteoblast MC3T3-E1 morphology revealed the colonization of pores in dentine bioapatites and an aligned cell growth in the oriented enameloid crystals. A higher proliferation (p < 0.01) was detected at up to 21 d in both the shark bioapatites and synthetic biphasic graft with respect to the bovine mineralized matrix. Finally, the great potential of porous biphasic dentine bioapatites enriched with Mg and the aligned fluorapatite crystals of enameloid bioapatites in promoting greater osteogenic activity was confirmed with a significantly increased ALP synthesis (p < 0.01) compared to the commercial grafts.

Strontium isotope age-dating of fossil shark tooth enameloid from the Upper Cretaceous Strata of Alabama and Mississippi, USA

Cretaceous strata in Alabama and Mississippi (USA) represent one of the most complete records of shallow marine deposition worldwide for the Upper Cretaceous. The age assignment of these strata in the eastern Gulf Coastal Plain is difficult due to the comparative lack of radiometrically datable beds and sometimes conflicting results of biostratigraphy using different taxonomic groups. Numerical age dating using strontium isotope ratios (87Sr/86Sr) preserved in diagenetically resistant fossil shark tooth enameloid had been proposed by previous researchers as a solution to dating some geologic units. Here we apply this methodology to the whole Upper Cretaceous, using teeth of two fossil shark genera (Scapanorhynchus and Squalicorax) collected from variable facies. Shark teeth collected from a bentonite mine in Monroe County, Mississippi, were also analyzed and compared with the radiometric date of the bentonite layer. Results indicate a strong correlation between stratigraphic position of the fossil teeth and numerical age determination based on 87Sr/86Sr content. Furthermore, this method is equally effective for both of the fossil shark genera analyzed in the study. Because of the nearly uniform distribution of strontium in ocean water, numerical age dating using strontium isotope ratios preserved in fossil shark tooth enameloid can be a useful method to employ in the correlation of marine geological strata on both regional and global scales.

Gradual changes in upwelled seawater conditions (redox, pH) from the late Cretaceous through early Paleogene at the northwest coast of Africa: Negative Ce anomaly trend recorded in fossil bio-apatite

Marine vertebrate fossils from late Cretaceous–early Paleogene shallow marine phosphorites of Morocco were investigated for their trace element compositions and neodymium isotopic ratios in order to constrain marine and depositional conditions. The various analyzed fossils are separated into two groups with different geochemical compositions: (1) shark tooth enameloid with generally lower Cu, Ba, rare earth elements (REEs) and U, and higher Na, Zn and Sr concentrations than (2) dentine, bone, and coprolites. These differences are related to the originally different structures of these fossils.All the fossils revealed very similar shale-normalized REE patterns, with negative Ce anomaly and heavy REE enrichment mimicking the REE pattern of modern oxic-seawater. The results therefore suggest REE uptake by the fossils from an early diagenetic pore fluid that was dominated by seawater. Importantly, the Ce anomaly does not vary among the different types of fossil remains and the values are very similar in a given layer. Moreover, a step-wise shift towards lower Ce/Ce* values from older to younger beds is apparent. Nd isotope analyses across the phosphorite succession yielded minor variation with an average εNd(t) value of −6.2±0.4 (n=12) indicating no major changes in the REE source during the studied interval. The relatively radiogenic values reflect Tethyan connection and/or important contribution of a mafic Nd source from weathering in the region.The age-related trend in the Ce anomaly is interpreted to be due to changes in redox and/or pH conditions of the upwelling seawater on the shallow shelf areas. The driving force of these changes was probably an extended global oceanic circulation, especially enhanced connection and water exchange between the North and South Atlantic.

Structure and mechanical properties of normal and anomalous teeth in the sand tiger shark Carcharias taurus.

This study reports and characterises an anomalous condition in teeth of captive sand tiger sharks Carcharias taurus . Abnormal shed teeth from captive sand tiger sharks which were soft to the touch were structurally and chemically characterised and compared to the structure and composition of normal teeth of the species. Normal specimens exhibited the expected tooth morphology with a well-developed tooth crown consisting of a central cusp and two lateral cusplets with smooth and sharp margins, while in the anomalous specimens the crown height was much reduced and the overall shape did not follow this pattern. Lateral cusplets were also considerably reduced in size and with blunt margins. Scanning electron images showed a distinct absence of crystalline structures in the anomalous specimens. Raman microscopy analysis confirmed the low volume of fluoroapatite in the outer layers of the anomalous teeth, while the composition of the inner layers corresponding to dentine was comparable to the normal tooth specimens. Nanoindentation-derived mechanical properties showed significant differences between the anomalous and normal teeth. The mean values for enameloid elastic modulus and hardness of all three normal teeth were 75.92±3.4 GPa and 3.27±0.41 GPa, respectively. On the other hand, mean values of elastic modulus and hardness for anomalous teeth were 7.81±3.27 GPa and 0.39±0.25 GPa, respectively. However, mechanical property values of the dentine of normal and anomalous teeth were similar. The mean values of dentine elastic modulus and hardness of the normal teeth were 25.66±2.14 GPa and 0.89±0.01 GPa, respectively, while mean values for the anomalous teeth were 25.34±1.54 GPa and 0.83±0.03 GPa. Although the morphological, mechanical and chemical differences between the normal and anomalous teeth are quite evident, establishing the causes of this condition are not possible at this stage.

Chondrichthyan tooth enameloid: past, present, and future

Enameloid is a hard mineralized tissue covering chondrichthyan and actinopterygian teeth. Over the past 40 years, it has been extensively studied in various extinct and extant sharks, leading to the broad use of microstructural characters to differentiate between hybodont and neoselachian teeth. However, the chondrichthyan taxic diversity is disproportionately high compared to the number of taxa explored for enameloid microstructure, and the generalization of these few observations to the whole group is problematic. Indeed, many other groups, in particular modern rays and skates, have been completely overlooked, and almost nothing is known about their tooth histology. Furthermore, the recent discovery of typical neoselachian character in cladodontomorph sharks teeth clearly indicates that we have had an over-simplified perception of the chondrichthyan enameloid distribution, which put into question the previously proposed evolutive scenarios dealing whith this tissue. We propose a brief historical overview of the study and understanding of chondrichthyan enameloid diversity and briefly discuss preparation issues encountered when dealing with the study of chondrichthyan hypermineralized tissues. Then, the variation of enameloid microstructures encountered in ctenacanthiforms, hybodonts, selachimorphs, and batomorphs is explored, summarized, and discussed. Although the full extent of the diversity and variability of the enameloid microstructure in many of these groups and others remains to be fully determined, we are able to show that most possess a much more complex enameloid microstructure than expected, and propose a revised and more fitting chondrichthyan enameloid terminology, based on the recognition of two main units: an external Single Crystallite Enameloid (SCE) and an internal Bundled Crystallite Enameloid (BCE). Our study reveals new insights in the understanding of character distribution among batomorphs and sets a framework for tackling global chondrichthyan tooth enameloid evolution. © 2015 The Linnean Society of London

The enameloid microstructure of euselachian (Chondrichthyes) scales

The enameloid microstucture of chondrichthyan teeth has been studied for decades and it has proven to be a useful taxonomic tool. Changes in enameloid organization have been related to the emergence of new trophic strategies and Mesozoic radiation of the neoselachian crown group. However, in contrast to the abundance of these data on tooth enameloid, descriptions of chondrichthyan scale enameloid are almost nonexistent. The topology and microstructure of scale enameloid in particular euselachian groups: fossil Mesozoic Hybodontiformes and living neoselachians, including batoids and sharks, are described. It is shown that a thick layer of single crystallite enameloid (SCE) covers all studied scales. Although the enameloid of scales clearly does not reach high levels of microstructural differentiation present in the dental enameloid of some neoselachians, we found some degree of organization, such as oriented crystallites, differentiation into sublayers, and the presence of poorly structured sets of densely arranged parallel crystallites. As scales lack feeding functions of teeth, we suggest that the emergence of microstructural organization/differentiation of chondrichthyan enameloid can be understood as consequence of a self-organizing process rather than adaptive pressure.

Ultrastructural organization and micromechanical properties of shark tooth enameloid

The outer part of shark teeth is formed by the hard and mineral-rich enameloid that has excellent mechanical properties, which makes it a very interesting model system for the development of new bio-inspired dental materials. We characterized the microstructure, chemical composition and resulting local mechanical properties of the enameloid from teeth of Isurus oxyrinchus (shortfin mako shark) by performing an in-depth analysis using various high-resolution analytical techniques, including scanning electron microscopy, qualitative energy-dispersive X-ray spectroscopy and nanoindentation. Shark tooth enameloid reveals an intricate hierarchical arrangement of thin (50–80nm) and long (>1μm) crystallites of fluoroapatite with a high degree of structural anisotropy, which leads to exceptional mechanical properties. Both stiffness and hardness are surprisingly homogeneous in the shiny layer as well as in the enameloid: although both tooth phases differ in structure and composition, they show almost no orientation dependence with respect to the loading direction of the enameloid crystallites. The results were used to determine the structural hierarchy of shark teeth, which can be used as a base for establishing design criteria for synthetic bio-inspired and biomimetic dental composites.