Development and Evolution of Dentition Pattern and Tooth Order in the Skates And Rays (Batoidea; Chondrichthyes)

Shark and ray (elasmobranch) dentitions are well known for their multiple generations of teeth, with isolated teeth being common in the fossil record. However, how the diverse dentitions characteristic of elasmobranchs form is still poorly understood. Data on the development and maintenance of the dental patterning in this major vertebrate group will allow comparisons to other morphologically diverse taxa, including the bony fishes, in order to identify shared pattern characters for the vertebrate dentition as a whole. Data is especially lacking from the Batoidea (skates and rays), hence our objective is to compile data on embryonic and adult batoid tooth development contributing to ordering of the dentition, from cleared and stained specimens and micro-CT scans, with 3D rendered models. We selected species (adult and embryonic) spanning phylogenetically significant batoid clades, such that our observations may raise questions about relationships within the batoids, particularly with respect to current molecular-based analyses. We include developmental data from embryos of recent model organisms Leucoraja erinacea and Raja clavata to evaluate the earliest establishment of the dentition. Characters of the batoid dentition investigated include alternate addition of teeth as offset successional tooth rows (versus single separate files), presence of a symphyseal initiator region (symphyseal tooth present, or absent, but with two parasymphyseal teeth) and a restriction to tooth addition along each jaw reducing the number of tooth families, relative to addition of successor teeth within each family. Our ultimate aim is to understand the shared characters of the batoids, and whether or not these dental characters are shared more broadly within elasmobranchs, by comparing these to dentitions in shark outgroups. These developmental morphological analyses will provide a solid basis to better understand dental evolution in these important vertebrate groups as well as the general plesiomorphic vertebrate dental condition.     

Tooth and consequences: Heterodonty and dental replacement in piranhas and pacus (Serrasalmidae)

Tooth replacement in piranhas is unusual: all teeth on one side of the head are lost as a unit, then replaced simultaneously. We used histology and microCT to examine tooth‐replacement modes across carnivorous piranhas and their herbivorous pacu cousins (Serrasalmidae) and then mapped replacement patterns onto a molecular phylogeny. Pacu teeth develop and are replaced in a manner like piranhas. For serrasalmids, unilateral tooth replacement is not an “all or nothing” phenomenon; we demonstrate that both sides of the jaws have developing tooth rows within them, albeit with one side more mineralized than the other. All serrasalmids (except one) share unilateral tooth replacement, so this is not an adaptation for carnivory. All serrasalmids have interlocking teeth; piranhas interdigitate lateral tooth cusps with adjacent teeth, forming a singular saw‐like blade, whereas lateral cusps in pacus clasp together. For serrasalmids to have an interlocking dentition, their teeth need to develop and erupt at the same time. We propose that interlocking mechanisms prevent tooth loss and ensure continued functionality of the feeding apparatus. Serrasalmid dentitions are ubiquitously heterodont, having incisiform and molariform dentitions reminiscent of mammals. Finally, we propose that simultaneous tooth replacement be considered as a synapomorphy for the family.     

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.     

Structure, attachment, replacement and growth of teeth in bluefish, Pomatomus saltatrix (), a teleost with deeply socketed teeth

Tooth replacement poses many questions about development, pattern formation, tooth attachment mechanisms, functional morphology and the evolution of vertebrate dentitions. Although most vertebrate species have polyphyodont dentitions, detailed knowledge of tooth structure and replacement is poor for most groups, particularly actinopterygians. We examined the oral dentition of the bluefish, Pomatomus saltatrix, a pelagic and coastal marine predator, using a sample of 50 individuals. The oral teeth are located on the dentary and premaxillary bones, and we scored each tooth locus in the dentary and premaxillary bones using a four-part functional classification: absent (A), incoming (I), functional (F=fully ankylosed) or eroding (E). The homodont oral teeth of Pomatomus are sharp, deeply socketed and firmly ankylosed to the bone of attachment. Replacement is intraosseus and occurs in alternate tooth loci with long waves of replacement passing from rear to front. The much higher percentage of functional as opposed to eroding teeth suggests that replacement rates are low but that individual teeth are quickly lost once erosion begins. Tooth number increases ontogenetically, ranging from 15–31 dentary teeth and 15–39 premaxillary teeth in the sample studied. Teeth increase in size with every replacement cycle. Remodeling of the attachment bone occurs continuously to accommodate growth. New tooth germs originate from a discontinuous dental lamina and migrate from the lingual (dentary) or labial (premaxillary) epithelium through pores in the bone of attachment into the resorption spaces beneath the existing teeth. Pomatomus shares unique aspects of tooth replacement with barracudas and other scombroids and this supports the interpretation that Pomatomus is more closely related to scombroids than to carangoids.     

Tooth development and replacement in the Atlantic Cutlassfish,Trichiurus lepturus,with comparisons to other Scombroidei

Atlantic Cutlassfish, Trichiurus lepturus, have large, barbed, premaxillary and dentary fangs, and sharp dagger-shaped teeth in their oral jaws. Functional teeth firmly ankylose to the dentigerous bones. We used dry skeletons, histology, SEM, and micro-CT scanning to study 92 specimens of T. lepturus from the western North Atlantic to describe its dentition and tooth replacement. We identified three modes of intraosseous tooth replacement in T. lepturus depending on the location of the tooth in the jaw. Mode 1 relates to replacement of premaxillary fangs, in which new tooth germs enter the lingual surface of the premaxilla, develop horizontally, and rotate into position. We suggest that growth of large fangs in the premaxilla is accommodated by this horizontal development. Mode 2 occurs for dentary fangs: new tooth germs enter the labial surface of the dentary, develop vertically, and erupt into position. Mode 3 describes replacement of lateral teeth, in which new tooth germs enter a trench along the crest of the dentigerous bone, develop vertically, and erupt into position. Such distinct modes of tooth replacement in a teleostean species are unknown. We compared modes of replacement in T. lepturus to 20 species of scombroids to explore the phylogenetic distribution of these three replacement modes. Alternate tooth replacement (in which new teeth erupt between two functional teeth), ankylosis, and intraosseous tooth development are plesiomorphic to Bluefish + other Scombroidei. Our study highlights the complexity and variability of intraosseous tooth replacement. Within tooth replacement systems, key variables include sites of formation of tooth germs, points of entry of tooth germs into dentigerous bones, coupling of tooth germ migration and bone erosion, whether teeth develop horizontally or immediately beneath the tooth to be replaced, and how tooth eruption and ankylosis occur. Developmentally different tooth replacement processes can yield remarkably similar dentitions.     

General morphology and ultrastructure of the radula of Testudinalia testudinalis (O. F. Müller, 1776) (Patellogastropoda,Gastropoda)

The radular morphology of the patellid species Testudinalia testudinalis (O. F. Müller, 1776) from the White Sea was studied using light, electron, and confocal microscopy. The radula is of the docoglossan type with four teeth per row and consisting of six zones. We characterize teeth formation in T. testidinalis as follows: one tooth is formed by numerous and extremely narrow odontoblasts through apocrine secretion; this initially formed tooth consists of numerous vesicles; the synthetic apparatus of the odontoblasts is localized in the apical and central parts of the cells throughout the cytoplasm and is penetrated by microtubules which are involved in the transport of the synthesized products to the apical part of the odontoblast; the newly formed teeth consist of unpolymerized chitin. Mitotic activity is located in the lateral parts of the formation zone. The first four rows contain an irregular arrangement of teeth, but the radular teeth are regularly arranged after the fifth row. The irregularly arranged teeth early on could be a consequence of the asynchronous formation of teeth and the distance between the odontoblasts and the membranoblasts. The morphological data obtained significantly expands our knowledge of the morphological diversity of the radula formation in Gastropoda.     

Marginal tooth-bearing bones in the lower jaw of the recent australian lungfish,Neoceratodus forsteri (Osteichthyes,Dipnoi)

Developmental studies of the Recent Australian lungfish, Neoceratodus forsteri, show that this species has two sets of functional tooth-bearing bones in the lower jaw of young hatchlings. These coincide with an early stage in the life history when the fish is strictly carnivorous. In N. forsteri, a paired tooth-bearing dentary and an unpaired symphyseal bone and tooth develop slightly later than the permanent vomerine, prearticular, and pterygopalatine tooth plates, which appear at stage 44 of development, and erupt with the permanent dentition between stages 46 and 48, when the hatchling first starts to feed on small aquatic invertebrates. At these stages of development, all of the teeth are long, sharp, and conical and help to retain prey items in the mouth. Disappearance of the transient dentition coincides with complete eruption of the permanent tooth plates and precedes the change to an omnivorous diet. Existence of a transient marginal dentition in this species of lungfish suggests that the presence of an apparently similar marginal dentition in adults of many species of Palaeozoic dipnoans should be considered in phylogenetic analyses of genera within the group, and when analysing the relationships of dipnoans with other primitive animals. © 1995 Wiley-Liss, Inc.     

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.     

Squatiniformes (Chondrichthyes,Neoselachii) from the Late Cretaceous of southern England and northern France with redescription of the holotype of Squatina cranei Woodward, 1888

Abstract: Bulk sampling of phosphate‐rich horizons within the Late Cretaceous of the Anglo‐Paris Basin yielded numerous teeth of members of the Squatiniformes. Along with isolated tooth remains, two museum specimens comprising partial articulated encoskeletal remains including the holotype of the species Squatina cranei Woodward, 1888a are described, and a new subgenus Cretascyllium is proposed for species of the genus Squatina with high degree of heterodonty and triangular anterior teeth. The species Squatina (Cretascyllium) cranei comb. nov. and Squatina (Cretascyllium) hassei comb. nov. are referred to this subgenus. The genus Parasquatina Herman, 1982 previously erected on a single tooth is valid, and two new species P. justinensis sp. nov. and P. jarvisi sp. nov. are described along with a third taxon Parasquatina sp. An enigmatic tooth referred to ?Neoselachii incertae sedis is also reported. The palaeoecology of these taxa is discussed.     

Ectoderm,endoderm, and the evolution of heterodont dentitions

Mammalian dentitions consist of different shapes/types of teeth that are positioned in different regions of the jaw (heterodont) whereas in many fish and reptiles all teeth are of similar type (homodont). The process by which heterodont dentitions have evolved in mammals is not understood. In many teleosts teeth develop in the pharynx from endoderm (endodermal teeth), whereas mammalian teeth develop from the oral ectoderm indicating that teeth can develop (and thus possibly evolve) via different mechanisms. In this article, we compare the molecular characteristics of pharyngeal/foregut endoderm with the molecular characteristics of oral ectoderm during mouse development. The expression domains of Claudin6, Hnf3β, α‐fetoprotein, Rbm35a, and Sox2 in the embryonic endoderm have boundaries overlapping the molar tooth‐forming region, but not the incisor region in the oral ectoderm. These results suggest that molar teeth (but not incisors) develop from epithelium that shares molecular characteristics with pharyngeal endoderm. This opens the possibility that the two different theories proposed for the evolution of teeth may both be correct. Multicuspid (eg. molars) having evolved from the externalization of endodermal teeth into the oral cavity and monocuspid (eg. incisors) having evolved from internalization of ectodermal armour odontodes of ancient fishes. The two different mechanisms of tooth development may have provided the developmental and genetic diversity on which evolution has acted to produce heterodont dentitions in mammals. genesis 48:382–389, 2010. © 2010 Wiley‐Liss, Inc.     

Making teeth to order: conserved genes reveal an ancient molecular pattern in paddlefish (Actinopterygii)

Ray-finned fishes (Actinopterygii) are the dominant vertebrate group today (+30 000 species, predominantly teleosts), with great morphological diversity, including their dentitions. How dental morphological variation evolved is best addressed by considering a range of taxa across actinopterygian phylogeny; here we examine the dentition of Polyodon spathula (American paddlefish), assigned to the basal group Acipenseriformes. Although teeth are present and functional in young individuals of Polyodon, they are completely absent in adults. Our current understanding of developmental genes operating in the dentition is primarily restricted to teleosts; we show that shh and bmp4, as highly conserved epithelial and mesenchymal genes for gnathostome tooth development, are similarly expressed at Polyodon tooth loci, thus extending this conserved developmental pattern within the Actinopterygii. These genes map spatio-temporal tooth initiation in Polyodon larvae and provide new data in both oral and pharyngeal tooth sites. Variation in cellular intensity of shh maps timing of tooth morphogenesis, revealing a second odontogenic wave as alternate sites within tooth rows, a dental pattern also present in more derived actinopterygians. Developmental timing for each tooth field in Polyodon follows a gradient, from rostral to caudal and ventral to dorsal, repeated during subsequent loss of teeth. The transitory Polyodon dentition is modified by cessation of tooth addition and loss. As such, Polyodon represents a basal actinopterygian model for the evolution of developmental novelty: initial conservation, followed by tooth loss, accommodating the adult trophic modification to filter-feeding.     

Redescription of a rare bothid,Asterorhombus bleekeri (Macleay), and description of a new species ofAsterorhombus from northwestern Australia (Teleostei: Pleuronectiformes)

A rare Australian bothid flounder.Asterorhombus bleekeri (Macleay), is redescribed from the holotype and ten additional specimens from the east coast of Qeensland, Gulf of Carpentaria and Rowly Shoal (Western Australia). The species is transferred fromArnoglossus Bleeker toAsterorhombus Tanaka because of the lack of obvious sexual dimorphism in the interorbital width and pectoral fin length, the lack of rostral and orbital spines, the yellow-white blind side body coloration, and the deeply cleft parhypural and hypural plates. The definition ofAsterorhombus was emended as follows: the first dorsal fin ray continuous with or separated from remaining fin rays and gill rakers slender or stubby, with or without serrations.Asterorhombus osculus sp. nov., formerly briefly described in the literature as unidentified species ofEngyprosopon, was described from eight specimens from the northwestern coast of Australia. The new species is most similar toA. bleekeri in lacking sexual dimorphism, and having the caudal skeleton with deep clefts, two or three rows of teeth on the upper jaw and a pair of conspicuous black spots on the caudal fin, in addition to a similar general appearance, but is distinguished from the latter by shorter gill rakers, a very small mouth and feebly ctenoid scales on the ocular side. Both species clearly differed fromA. intermedius andA. fijiensis in having two (or three) rows of teeth on the upper jaws, slender gill rakers without serrations, first dorsal fin ray continuous with the other fin rays, and a pair of conspicuous black spots on the caudal fin.     

Chondrichthyans from a Cenomanian (Late Cretaceous) bonebed,Saskatchewan, Canada

Abstract: Acid preparation of samples of a bonebed from the Cenomanian of central Canada yielded several thousand well‐preserved chondrichthyan teeth, in addition to numerous other vertebrate remains. Teeth and other remains of one species of chimaeroid, one species of hybodont shark, three species of Ptychodus, 10 species of neoselachian sharks and two species of batoid were recorded. The family Archaeolamnidae fam. nov., genera Meristodonoides gen. nov. and Telodontaspis gen. nov. and species Ptychodus rhombodus sp. nov., Telodontaspis agassizensis gen et sp. nov., Eostriatolamia paucicorrugata sp. nov., Roulletia canadensis sp. nov., Cretorectolobus robustus sp. nov. and Orectoloboides angulatus sp. nov. are described. Status of the genus Palaeoanacorax and the species Cretoxyrhina denticulata, Squalicorax curvatus and ‘Rhinobatos’incertus are discussed, and reconstructed dentitions of Archaeolamna and Roulletia presented. The fauna is of low diversity and dominated by active hunters, with many species apparently endemic to the northern Western Interior Seaway.     

Revision of the genusgirella (girellidae) from east asia

The taxonomy of three species ofGirella from East Asian waters,G. punctata Gray, 1835,G. leonina (Richardson, 1846) andG. mezina Jordan & Starks, 1907, is reviewed and intraspecific (individual and ontogenic) variations detailed.Girella mezina is characterized by a very wide mouth and thick upper lip, the soft-rayed portion of the anal fin high and round, dorsal profile of the head abruptly slanting in front of the eyes in adults, a transverse yellow band on the body in life, and 3–4 rows of teeth along the outer jaw margins, the central cusp of each tooth being wider than the lateral cusps in adults.Girella punctata is characterized by usually 2 rows of teeth along the outer jaw margins, usually 7 transverse series of scales between the lateral line and median spinous portion of the dorsal fin (TRac), and usually 52–55 pored lateral line scales (LLp). The species is variable in body and caudal fin shape, extent of squamation on the opercular region, and number and position of dark spots on the scales. It also exhibits ontogenetic variation in the number of tooth-rows.Girella leonina is characterized by a conspicuously black opercular flap, essentially a single row of teeth along the outer jaw margins, usually 10–11 TRac and usually 59–64 LLp. Intraspecific variations were evident in the mouth position, scale condition and body color after death, and in the number of pores of cephalic lateral line canals. The holotype ofG. punctata, previously known only from a figure, is described for the first time.Girella melanichthys is synonymized underG. punctata with a lectotype designated for the former.     

3D morphology of pharyngeal dentition of the genus Capoeta (Cyprinidae): Implications for taxonomy and phylogeny

Capoeta is a herbivorous cyprinid fish genus, widely distributed in water bodies of Western Asia. Recent species show a distinct biogeographic pattern with endemic distribution in large fluvial drainage basins. As other cyprinids, the species of this genus are characterized by the presence of the pharyngeal bone with pharyngeal teeth. Despite this, the detailed morphology of the pharyngeal teeth, its interspecific and topologic variations, and the importance for taxonomy and phylogeny of the genus Capoeta are still not established. For the first time, a detailed comprehensive study of the pharyngeal dentition of 10 Capoeta species has been provided. The morphologic study of the pharyngeal dentition bases on the 3D microtomography and follows the purpose to evaluate the potential taxonomic and phylogenetic signals of these elements, as well as to study interspecific and topologic variations of the pharyngeal teeth. In this study, we propose a new methodology to categorize the studied pharyngeal teeth in 18 shape classes. The results of this study show that the detailed 3D morphology of the pharyngeal teeth is a useful tool for the identification of isolated teeth at the generic and/or specific level and that in certain cases, the tooth position in the teeth rows can be identified. Additionally, the preliminary analysis shows that the morphology of the pharyngeal teeth provides a potential phylogenetic signal. Both these patterns are very important for the taxonomy of cyprinid fishes and especially can be applied to fossil records.     

The oral apparatus of tadpoles of Rana dalmatina, Bombina variegata, Bufo bufo, and Bufo viridis (Anura)

The morphology and development of the larval oral apparatus of Rana dalmatina, Bombina variegata, Bufo bufo, and Bufo viridis are described and compared using scanning electron microscopy. The species show different arrangements of the mouthparts. The small oral apparatus of R. dalmatina larvae has three labial tooth rows on the upper labium, while there are four tooth rows on the lower labium with a medial gap in row proximal to the mouth. The margins of the oral apparatus are defined by papillae that encircle the lower labium. B. variegata tadpoles have two upper labial tooth rows and three lower labial tooth rows that are uninterrupted, unlike the ones of R. dalmatina. The mouth is encircled by papillae that are larger than those of R. dalmatina. The oral discs of tadpoles of both B. bufo and B. viridis are similar. They are defined by two upper labial tooth rows (the second of which is interrupted by a medial gap) and by three lower tooth rows that differ in lengths in the two Bufo species. Both species develop papillae on the mouth angles and in two rows on the upper labium. Some morphological differences among the oral discs of R. dalmatina, B. variegata, B. bufo, and B. viridis tadpoles can be attributed to phylogenetic differences, but most can be related to their varying feeding habits and/or to their dietary specializations.     

New species of the genus <Emphasis Type="Italic">Bathycongrus—B. parviporus</Emphasis> (Congridae,Anguilliformes)—from waters of Central Vietnam (Nha Trang and Van Phong Bays)

Bathycongrus parviporus sp. nova is described from specimens collected in the South China Sea, in the coastal waters of central Vietnam. By its characters (short snout, elongate-oval vomerine tooth patch with numerous small teeth of approximately the same size, slender relatively short tail, and residual leptocephalic pigmentation as a series of small melanophores just below lateral line in adults), the species belongs to the group of species of this genus represented by B. bleekeri, B. trimaculatus, and B. unimaculatus in the western part of the Pacific Ocean and B. dubius in the western Atlantic Ocean but differs in small infraorbital pores which are enlarged in all known species of the genus Bathycongrus. By the number of preanal pores, the new species is similar to B. bleekeri from Philippine waters but differs from it in a higher number of vertebrae (120–122), in more numerous rows of teeth on the premaxilloethmoid, in more number of branchiostegal rays (9–10), a shorter head, and in some other proportions of head and body. Topography of canals and pores of the cephalic seismosensory system of B. parviporus sp. nova is described. Morphometric and osteological characters for placing this species in the genus Bathycongrus are noted.     

Mandibular ontogeny in the miocene great apeDryopithecus

The type mandible of Dryopithecus fontani,Lartet 1856, has been discovered to be not fully adult. Its development corresponds in dental age to that of a 6-to 8-year-old chimpanzee. Because of its immaturity, a number of seemingly distinctive features of this mandible (some of which resemble hominids) would have been lost with full adulthood. Closed tooth rows, a recurved canine, and a vertical ascending mandibular ramus are related to the age of the specimen. They therefore do not foreshadow hominid characteristics. It is stressed that consideration of individual age is an important factor in interpreting the dentitions of fossil and extant hominoids.     

Two new species of Leporinus (Characiformes: Anostomidae) from the Brazilian Amazon,and redescription of Leporinus striatus Kner 1858

Two new species of Leporinus are described from tributaries of the Rio Amazonas in Brazil. One species is known from the Jari and Tapajós River basins, and is identified on the basis of a gas bladder reduced in size, a dark midlateral stripe on the body, dark transverse bars on the dorsum, a subinferior mouth, three teeth on the premaxilla, four teeth on the dentary and 16 scale rows around the caudal peduncle. The second new species is known from the Tocantins, Xingu and Tapajós River basins, and is identified on the basis of three dark longitudinal stripes on the body, a subinferior mouth, three teeth on the premaxilla, four teeth on the dentary and 12 scale rows around the caudal peduncle. In addition, Leporinus striatus is redescribed based on type and additional specimens from the Río de La Plata, Amazonas, Orinoco, Atrato, Magdalena and Sinu River basins. Leporinus striatus is identified on the basis of four dark longitudinal stripes on the body, a subterminal mouth, three teeth on the premaxilla, four teeth on the dentary and 16 scale rows around the caudal peduncle.