The dentition and dental tissues of the agamid lizard, Uromastyx

The dentition of Uromastyx hardwicki was examined in a series of carefully prepared dry skulls and was found to be very different from that of other agamid lizards. The anatomy of the dentition undergoes great changes from the time of hatching to advanced age, but no evidence of tooth replacement could be found. Extension of the tooth rows by addition of larger teeth posteriorly, together with elongation of the premaxilla, and a characteristic pattern of wear are responsible for the condition seen in aged specimens.
The structure of the dental tissues was investigated by means of a variety of histological techniques including scanning electron microscopy and it is established that the enamel has prismatic structure like that of mammalian enamel. The mode of formation of enamel with and without prisms is described and the occurrence and significance of prismatic structure in reptilian dental enamel discussed.     

Saurian malaria in Kenya: epidemiological features of malarial infections in lizard populations of the West Pokot District

During investigations into the prevalence of malarial parasites among lizards in the West Pokot District in Kenya, 179 lizards comprising eight species were caught. Examination of the Giemsa-stained smears made from their blood showed that 34 lizards were infected with Plasmodium species. Fifteen lizards were infected with a single species of Plasmodium and 19 carried multiple infections, the maximum, in four lizards, was four species. There were 19 combinations of parasite infections. Seventeen Plasmodium species were identified, the commonest being P. icipeensis. Only two of the eight lizard species were infected: the skink Mabuya striata and the agamid Agama agama. Eight of the Plasmodium species infected both; another eight species infected M. striata only but three of these have been described from different lizard families elsewhere in Africa. P. robinsoni infected A. agama only, although it was first described from another lizard family in another part of Africa. The epidemiological significance of these results is discussed.     

Fight versus flight: Body temperature influences defensive responses of lizards

In laboratory studies we determined that the defensive responses used by two agamid lizards, Agama savignyi and A. pallida, change as a function of body temperature. At high body temperatures, these lizards flee rapidly from predators. At lower body temperatures, which reduce sprint speed, the lizards rarely run but instead hold their ground and attack aggressively. This temperature-dependent switch in defensive behaviour may have evolved because cold lizards that live in open habitats would have little chance of outrunning predators. Defensive behaviours of animals may in general be sensitive to physiological variables that influence locomotor performance.     

Ontogenetic transition from unicuspid to multicuspid oral dentition in a teleost fish: Astyanax mexicanus, the Mexican tetra (Ostariophysi: Characidae)

Teleost fishes display a remarkable diversity of adult dentitions; this diversity is all the more remarkable in light of the uniformity of first-generation dentitions. Few studies have quantitatively documented the transition between generalized first-generation dentitions and specialized adult dentitions in teleosts. We investigated this transition in the Mexican tetra, Astyanax mexicanus (Characidae), by measuring aspects of the dentition in an ontogenetic series of individuals from embryos to 160 days old, in addition to adults of unknown age. The first-generation dentition and its immediate successors consist of small, unicuspid teeth that develop extraosseously. Multicuspid teeth first appear during the second tooth replacement event, and are derived from single tooth germs, rather than from the fusion of multiple conical tooth germs. We document that the transition from unicuspid to multicuspid teeth corresponds to a change in the location of developing tooth germs (from extraosseous to intraosseous) and in patterns of tooth replacement (from haphazard to simultaneous within a jaw quadrant). In addition, while the size of the largest teeth scales with positive allometry to fish size, the transition to multicuspid teeth is accompanied by an exceptionally large increase in tooth size.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 523–538.     

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.     

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.     

Numeric anomalies of teeth in concomitant hypodontia and hyperdontia

Variations in tooth number in children, each of whom had supernumerary teeth and agenesis of teeth, is described. Among the 11, seven had cleft lip and palate, and two had clefting syndromes; two children had dental anomalies only. Only children who had both supernumerary teeth and congenitally missing teeth outside the area of the cleft alveolus were included. Concomitant hypodontia and hyperdontia were observed in the same dentition in nine subjects, in the same jaw in eight subjects, and in the same jaw quadrant in only three subjects. Supernumerary teeth and agenesis of teeth were observed simultaneously more often in the permanent dentitions than in the deciduous dentitions or in both dentitions simultaneously. The overall number of supernumeraries was 10 in the deciduous dentition and 14 in the permanent dentition of the 11 subjects. The number of congenitally absent teeth was 14 in the deciduous dentition and 40 in the permanent dentition. The etiology of concomitant hypodontia and hyperdontia is difficult to explain. It may result from disturbances in migration, proliferation, and differentiation of neural crest cells or interactions between the epithelial and mesenchymal cells during the initiation of odontogenesis.     

Conserved deployment of genes during odontogenesis across osteichthyans

Odontogenesis has only been closely scrutinized at the molecular level in the mouse, an animal with an extremely restricted dentition of only two types and one set. However, within osteichthyans many species display complex and extensive dentitions, which questions the extent to which information from the mouse is applicable to all osteichthyans. We present novel comparative molecular and morphological data in the rainbow trout (Oncorhynchus mykiss) that show that three genes, essential for murine odontogenesis, follow identical spatial-temporal expression. Thus, at all tooth bud sites, epithelial genes Pitx-2 and Shh initiate the odontogenic cascade, resulting in dental mesenchymal Bmp-4 expression, importantly, including the previously unknown formation of replacement teeth. Significantly, this spatial-temporal sequence is the same for marginal and lingual dentitions, but we find notable differences regarding the deployment of Pitx-2 in the developing pharyngeal dentition. This difference may be highly significant in relation to the theory that dentitions may have evolved from pharyngeal tooth sets in jawless fishes. We have provided the first data on operational genes in tooth development to show that the same signalling genes choreograph this evolutionary stable event in fishes since the osteichthyan divergence 420 Myr ago, with the identical spatial-temporal expression as in mammals.     

The development and replacement of teeth in viviparous caecilians.

Tooth development and replacement in fetal and adult viviparous caecilians (Amphibia: Gymnophiona) are described and analyzed according to current theories of tooth succession. The fetal dentition differs from that of the adult in morphology, position, and function. Teeth are used by fetuses to scrape the oviducal epithelium, thus stimulating the secretion of a nutrient substance. Fetal dentitions vary in morphology and position in different species. The ontogeny of teeth of several species is described and the patterns of addition of loci and of replacement are analyzed. Loci are added both posteriorly along the jaw and between existing loci as the jaw grows prior to ossification; subsequently addition is restricted to the posterior part of the jaw. Tooth replacement is alternate. The several rows and patches of teeth are the result of retention of replacement series on the dentigerous elements. Tooth development and replacement in a series of juveniles and adults of different sizes in a single species are also considered. Post-fetal patterns of development and replacement are similar to those seen in larvae and adults of oviparous species. Variation in numbers of teeth and proportions of teeth at particular stages occurs ontogenetically and among individuals of the same size, though proportions occur in a similar pattern throughout the series. The general pattern of tooth replacement in fetuses and adults can be explained by either Edmund's Zahnreihen theory or by Osborn's Tooth Family theory, but replacement in fetal tooth patches and the fetal-adult dentitional transition are explained by neither.     

Oral food processing in two herbivorous lizards, Iguana iguana (Iguanidae) and Uromastix aegyptius (Agamidae).

The anatomy and function of the feeding apparatus in Iguana iguana and Uromastix aegyptius were studied by dissection, cinematic and cineradiographic techniques. The feeding behavior of these species differs from that of insectivorous lizards in the cropping action involves movement of both the upper jaw around the atlantooccipital joint and the lower jaw around the mandibular joint; and in Uromastix only, streptostylic movement of the quadrate. Often movements of the whole head play a supplementary role in the cropping action. In both species the feeding apparatus has been modified to facilitate cropping. In Iguana the pleurodont dentition is multicusped and laterally compressed. Each tooth forms a shearing blade whose function does not require contact with other teeth. In Uromastix the dentition is acrodont and the cheek teeth are massive and lack cusps. Occlusion is necessary for shearing plant material. The skull system of Uromastix also has a number of modified structures which allow protraction and retraction of the lower jaw to facilitate cropping while maintaining a gape equivalent to that in Iguana. It is suggested that the differences in the feeding apparatus between Iguana and Uromastix are attributable to differeces in the mode of tooth replacement and implantation.     

Geographic variation in Agama impalearis from Morocco: evidence for historical population vicariance and current climatic effects

Morphological and molecular surveys of within-island geographic variation in lizards have revealed patterns of geographic variation that reflect both population vicariance and in situ selection-mediated responses to the ecological heterogeneity of the islands. This study tested different models of differentiation in a continental species, the agamid Agama impalearis , in which spatial separation is much greater than that in island species. Patterns of among-site differentiation in morphology were described and a character-resampling technique used to investigate their robustness. Putative causes were evaluated by testing multivariate patterns of differentiation against models based on historical and present-day effects using matrix association randomization tests. This strongly suggested the action of both vicariance/secondary contact and current climatic conditions in shaping the patterns of morphological variation. The former appears to be the result of range contraction into refugia separated by the Atlas mountain range during glacial conditions, with subsequent secondary contact during warmer interglacial periods, Scalation showed a very clear association with geographic difference in thermal regime, even after the vicariance/gene-flow induced non-independence had been taken into account. This latter finding indicates that patterns observed in other lizards are quite general.     

Hinged teeth for hard-bodied prey: a case of convergent evolution between snakes and legless lizards

Savitzky (1981) described hinged teeth in several taxa of snakes, and interpreted this type of dentition as an adaptation to feeding on hard-bodied prey (scincid lizards). We tested this hypothesis by examining the dentition of insectivorous and saurophagous members of the Australian legless lizards, Pygopodidae. Insectivorous taxa ( Delma, Pygopus ) have peg-like pleurodont dentition, but the saurophagous Lialis has slender, recurved, sharply-pointed teeth, like those of many snakes. The teeth of Lialis are 'hinged' on their supporting bones: each tooth folds when pressure is applied to its anterior surface, but locks in an erect position when forced from behind, The tooth hinge is probably collagenous, and does not contain elastin. The presence of hinged teeth in Lids , which feeds predominately on scincid lizards, offers strong support for Savitzky's hypothesis.     

The deciduous dentition and dental replacement in the Eocene bat Palaeochiropteryx tupaiodon from Messel: The primitive condition and beginning of specialization of milk teeth among Chiroptera

The deciduous dentition and tooth replacement pattern of Palaeochiropteryx tupaiodon from the early Middle Eocene of Messel, near Frankfurt, Germany, are described. Ontogenetic states include fetuses to subadults. The posterior portion of the deciduous dentition (dP3-4) still shows the primitive eutherian condition of molarization, while the anterior part (dI-dC) was already engaged in the evolution of the highly derived condition found in living bats for clinging to the mother's fur. A styliform and sharp anterior dentition is considered a prerequisite in earliest chiropteran evolution. The greatly modified milk teeth of all living bats developed in different clades by parallel evolution under high selective pressure. The tiny and, at initial stages, poorly calcified teeth are substantiated by a newly developed microradiographic technique which is described in detail.     

Tooth replacement pattern of Coloborhynchus robustus (Pterosauria) from the Lower Cretaceous of Brazil

The well preserved anterior upper and lower jaw fragment of an adult specimen of Coloborhynchus robustus (Pterosauria: Ornithocheiridae), SMNK 2302 PAL, allowed investigations of the replacement pattern of the dentition macroscopically and by using CT scans. The quantification of the dentition by Zahnreihen, Z-Spacing, and replacement waves indicates a complex pattern of different replacement stages in which large gaps within the dentition were avoided. The specialized prey-catching apparatus of Coloborhynchus thus could retain its function even following tooth replacement. The replacement process in the specimen took about 2/3 of the total life-time of a tooth, and damaged teeth in the anterior jaw region may have been replaced more rapidly than posterior teeth. The distolingual replacement of the functional teeth delayed the time of their shedding in comparison with the circular resorption present in crocodiles. In contrast to these, the distolingual position of the replacement tooth did not decrease the biomechanical stability of the functional tooth, which can also be observed as a convergence in other thecodont dentitions, e.g., recent carnivore mammals. Teeth were shed when their replacement had reached about 60% of the full-grown height. A comparison of the observed pattern is constricted by the preservation and preparation of other specimens. Unfortunately, no known specimen in public collections reaches the quality of Coloborhynchus robustus, SMNK 2302 PAL, so that comparable patterns in other specimens are not likely to be detected.     

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.     

The histology of tooth formation in the Australian lungfish, Neoceratodus forsteri Krefft

The histology of developing toothplates of Neoceratodusforsteri from the time of first appearance of the tooth primordia to the adult condition has been investigated. The dentition develops by the formation of a shell of primary epithelial and mesenchymal matrices. Within the shell, secondary mesenchymal matrix and central material, both containing columns of tertiary matrix, are laid down. Primary epidielial matrix appears to contain collagen and is closely associated with the epithelium of the mouth. All other tooth tissues as well as the supporting bone develop in association with mesenchyme. Primary, secondary and tertiary mesenchymal matrices appear to contain collagen. Central dentine contains some fibres, possibly of reticulin or collagen, within a matrix of unknown composition.
The tooth is attached to the underlying bone by a pedestal of bone and this grows with the tooth material.
New tooth tissues are formed in the pulp cavity in layers below the older material, causing the toothplate to grow in every dimension as the animal grows.
An evolutionary pathway is suggested for lungfish with a dentition of cusps arranged in radiating ridges.     

Early development of rostrum saw-teeth in a fossil ray tests classical theories of the evolution of vertebrate dentitions

In classical theory, teeth of vertebrate dentitions evolved from co-option of external skin denticles into the oral cavity. This hypothesis predicts that ordered tooth arrangement and regulated replacement in the oral dentition were also derived from skin denticles. The fossil batoid ray Schizorhiza stromeri (Chondrichthyes; Cretaceous) provides a test of this theory. Schizorhiza preserves an extended cartilaginous rostrum with closely spaced, alternating saw-teeth, different from sawfish and sawsharks today. Multiple replacement teeth reveal unique new data from micro-CT scanning, showing how the ‘cone-in-cone’ series of ordered saw-teeth sets arrange themselves developmentally, to become enclosed by the roots of pre-existing saw-teeth. At the rostrum tip, newly developing saw-teeth are present, as mineralized crown tips within a vascular, cartilaginous furrow; these reorient via two 90° rotations then relocate laterally between previously formed roots. Saw-tooth replacement slows mid-rostrum where fewer saw-teeth are regenerated. These exceptional developmental data reveal regulated order for serial self-renewal, maintaining the saw edge with ever-increasing saw-tooth size. This mimics tooth replacement in chondrichthyans, but differs in the crown reorientation and their enclosure directly between roots of predecessor saw-teeth. Schizorhiza saw-tooth development is decoupled from the jaw teeth and their replacement, dependent on a dental lamina. This highly specialized rostral saw, derived from diversification of skin denticles, is distinct from the dentition and demonstrates the potential developmental plasticity of skin denticles.     

Early development and replacement of the stickleback dentition

Teeth have long served as a model system to study basic questions about vertebrate organogenesis, morphogenesis, and evolution. In nonmammalian vertebrates, teeth typically regenerate throughout adult life. Fish have evolved a tremendous diversity in dental patterning in both their oral and pharyngeal dentitions, offering numerous opportunities to study how morphology develops, regenerates, and evolves in different lineages. Threespine stickleback fish (Gasterosteus aculeatus) have emerged as a new system to study how morphology evolves, and provide a particularly powerful system to study the development and evolution of dental morphology. Here, we describe the oral and pharyngeal dentitions of stickleback fish, providing additional morphological, histological, and molecular evidence for homology of oral and pharyngeal teeth. Focusing on the ventral pharyngeal dentition in a dense developmental time course of lab‐reared fish, we describe the temporal and spatial consensus sequence of early tooth formation. Early in development, this sequence is highly stereotypical and consists of seventeen primary teeth forming the early tooth field, followed by the first tooth replacement event. Comparing this detailed morphological and ontogenetic sequence to that described in other fish reveals that major changes to how dental morphology arises and regenerates have evolved across different fish lineages. J. Morphol. 277:1072–1083, 2016. © 2016 Wiley Periodicals, Inc.     

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.