Feeding‐related characters in basal pterosaurs: implications for jaw mechanism,dental function and diet

?si, A. 2011: Feeding‐related characters in basal pterosaurs: implications for jaw mechanism, dental function and diet. Lethaia, Vol. 44, pp. 136–152. A comparative study of various feeding‐related features in basal pterosaurs reveals a significant change in feeding strategies during the early evolutionary history of the group. These features are related to the skull architecture (e.g. quadrate morphology and orientation, jaw joint), dentition (e.g. crown morphology, wear patterns), reconstructed adductor musculature and post‐cranium. The most basal pterosaurs (Preondactylus, dimorphodontids and anurognathids) were small‐bodied animals with a wingspan no greater than 1.5 m, a relatively short, lightly constructed skull, straight mandibles with a large gape, sharply pointed teeth and well‐developed external adductors. The absence of extended tooth wear excludes complex oral food processing and indicates that jaw closure was simply orthal. Features of these basal‐most forms indicate a predominantly insectivorous diet. Among stratigraphically older but more derived forms (Eudimorphodon, Carniadactylus, Caviramus) complex, multicuspid teeth allowed the consumption of a wider variety of prey via a more effective form of food processing. This is supported by heavy dental wear in all forms with multicuspid teeth. Typical piscivorous forms occurred no earlier than the Early Jurassic, and are characterized by widely spaced, enlarged procumbent teeth forming a fish grab and an anteriorly inclined quadrate that permitted only a relatively small gape. In addition, the skull became more elongate and body size increased. Besides the dominance of piscivory, dental morphology and the scarcity of tooth wear reflect accidental dental occlusion that could have been caused by the capturing or seasonal consumption of harder food items. □Basal pterosaurs, heterodonty, dental wear, insectivory, piscivory.     

Changes in orientation of attritional wear facets with implications for jaw motion in a mixed longitudinal sample of Propithecus edwardsi from Ranomafana National Park, Madagascar

In many mammalian species, the progressive wearing down of the teeth that occurs over an individual's lifetime has the potential to change dental function, jaw movements, or even feeding habits. The orientation of phase-I wear facets on molars reveals the direction of jaw movement during the power stroke of mastication. We investigated if and how molar wear facets change with increasing wear and/or age by examining a mixed longitudinal dataset of mandibular tooth molds from wild Propithecus edwardsi (N = 32 individuals, 86 samples). Measurements of the verticality of wear facets were obtained from three-dimensional digital models generated from μCT scans. Results show that verticality decreases over the lifetime of P. edwardsi, a change that implies an increasingly lateral translation of the jaw as the teeth move into occlusion. A more transverse phase-I power stroke supports the hypothesis that these animals chew to maximize longevity and functionality of their teeth, minimizing the "waste" of enamel, while maintaining sharp shearing crests. Results of this study indicate that wear facet verticality is more closely correlated with age than overall amount of tooth wear, measured as area of exposed dentin, suggesting that age-related changes in cranial morphology may be more responsible for adjustments in jaw motion over the lifetimes of Propithecus than wear-related changes inthe shape of occluding teeth. Finally, the rate of decrease in wear facet verticality with age is greater in males than in females suggesting differences in development and/or access to resources between the sexes in this species.     

Jaw movements from microwear on the molar teeth of the koala Phascolarctos cinereus

To study the direction of jaw movements in the koala from wear facets on the molar teeth by scanning electron microscopy, gold coated epoxy resin replicas from the right maxillary and mandibular tooth quadrants were examined from 12 koala skulls. The progressive development and location of facets, the orientation of striae on them and directional data were recorded and transferred from electron micrographs to superimposable transparencies.
Polished facets with laterally oriented striations developed on the cristids and cristae progressively into dentine, where Greaves' effect indicated that the direction of the chewing stroke was labiolingual. Polished and pitted facets, aligned and striated in the parasagittal plane, occurred on the smooth interactive enamel surfaces of maxillary and mandibular cusps.
Labiolingual transit of the crislids over the cristae, with a slight anteromedial shift, was inferred to be the predominant chewing stroke on the working side with no contralateral balancing contact. A propalinal isognathous movement in which successive cusps made contact was also deduced.
Previous concepts of koala chewing and tooth wear were confirmed and amplified, and these may have application to studies of extinct marsupial jaw mechanisms.     

Molar occlusion and jaw mechanics of the Eocene primate Adapis

Wear facets on molars of the Eocene primate Adapis magnus are described. Striations on these wear facets indicate three separate directions of mandibular movement during mastication. One direction corresponds to a first stage of mastication involving orthal retraction of the mandible. The remaining two directions correspond to buccal and lingual phases of a second stage of mastication involving a transverse movement of the mandible. The mechanics of jaw adduction are analysed for both the orthal retraction and transverse stages of mastication. During the orthal retraction stage the greatest component of bite force is provided by the temporalis muscles acting directly against the food with the mandible functioning as a link rather than as a lever. A geometrical argument suggests that during the transverse stage of mastication bite force is provided by the temporalis muscles of both sides, the ipsilateral medial and lateral pterygoid muscles, and the contralateral masseter muscle.     

Morphology and growth of lepidosirenid lungfish tooth plates (Pisces: Dipnoi)

The structure of the tooth plates of Protopterus and Lepidosiren was investigated to determine the causes and consequences of postlarval shape change. During growth, the basal area of the tooth plates increases, some cusps become more prominent, and shearing surfaces are sharpened. The jaw articulation restricts the range of movements of the lower jaw, and causes the tooth plates to occlude precisely; the resulting wear patterns are regular. The tooth plates are composed of enamel, trabecular dentine, and petrodentine. A petrodentine column forms the core of a tooth plate; it is flanked by trabecular dentine. Microhardness measurements show that trabecular dentine is comparable in hardness to mammalian dentine, whereas the petrodentine is comparable to enamel. The location and differential wear of these tissues produce the prominent cusps and self-sharpened blades of the adult tooth plates.     

Why fuse the mandibular symphysis? A comparative analysis

Fused symphyses, which evolved independently in several mammalian taxa, including anthropoids, are stiffer and stronger than unfused symphyses. This paper tests the hypothesis that orientations of tooth movements during occlusion are the primary basis for variations in symphyseal fusion. Mammals whose teeth have primarily dorsally oriented occlusal trajectories and/or rotate their mandibles during occlusion will not benefit from symphyseal fusion because it prevents independent mandibular movements and because unfused symphyses transfer dorsally oriented forces with equal efficiency; mammals with predominantly transverse power strokes are predicted to benefit from symphyseal fusion or greatly restricted mediolateral movement at the symphysis in order to increase force transfer efficiency across the symphysis in the transverse plane. These hypotheses are tested with comparative data on symphyseal and occlusal morphology in several mammals, and with kinematic and EMG analyses of mastication in opossums (Didelphis virginiana) and goats (Capra hircus) that are compared with published data on chewing in primates. Among mammals, symphyseal fusion or a morphology that greatly restricts movement correlates significantly with occlusal orientation: species with more transversely oriented occlusal planes tend to have fused symphyses. The ratio of working- to balancing-side adductor muscle force in goats and opossums is close to 1:1, as in macaques, but goats and opossums have mandibles that rotate independently during occlusion, and have predominantly vertically oriented tooth movements during the power stroke. Symphyseal fusion is therefore most likely an adaptation for increasing the efficiency of transfer of transversely oriented occlusal forces in mammals whose mandibles do not rotate independently during the power stroke.     

Tooth crown heights, tooth wear, sexual dimorphism and jaw growth in hominoids

The aim of this review is to bring together data that link tooth morphology with tooth function and tooth growth: We aim to show how the microanatomy of hominoid teeth is providing evidence about rates of tooth growth that are likely to be a consequence of both masticatory strategy and social behaviour. First, we present data about incisor and molar tooth wear in wild short chimpanzees that demonstrate how crown heights are likely to be related to relative tooth use in a broad sense. Following this we review recent studies that describe the microanatomy of hominoid tooth enamel and show how these studies are providing evidence about tooth crown formation times in hominoids, as well as improving estimates for the age at death of certain juvenile fossil hominids. Next, we outline what is known about the mechanisms of tooth growth in the sexually dimorphic canine teeth of chimpanzees and compare these patterns of growth with tooth growth patterns in the canines of three fossil hominids from Laetoli, Tanzania. Finally, we discuss how selection pressures that operate to increase or reduce the size of anterior teeth interact with jaw size. We argue that the space available to grow developing teeth in the mandibles of juvenile hominoids is determined by the growth patterns of the mandibles, which in turn reflect masticatory strategy. The consequences of selection pressure to grow large or small anterior teeth are likely to be reflected in the times at which these teeth are able to emerge into occlusion.     

Complexity of ruminant masticatory evolution

The evolution of robust jaws, hypsodont teeth, and large chewing muscles among grazing ruminants is a quintessential example of putative morphological adaptation. However, the degree of correlated evolution (i.e., to what extent the grazer feeding apparatus represents an evolutionary module), especially of soft and hard tissues, remains poorly understood. Recent generation of large datasets and phylogenetic information has made testing hypotheses of correlated evolution possible. We, therefore, test for correlated evolution among various traits of the ruminant masticatory apparatus including tooth crown height, jaw robustness, chewing muscle size, and characters of the molar occlusal surfaces, using phylogenetic and nonphylogenetic comparative methods as well as phylogenetic evolutionary model selection. We find that the large masseter muscles of grazing ruminants evolved with the inclusion of grass in the diet, an increase in the proportion of occlusal enamel bands oriented parallel to the chewing stroke, and possibly hypsodonty. We suggest that the masseter evolved under two evolutionary regimes: i) selection for higher masticatory forces during chewing and ii) flattening of the tooth profile, which resulted in reduced tooth guidance and, thus, a requirement for more chewing muscle activity during each chewing stroke, in agreement with previous research. The linear jaw metrics (depth of the mandibular angle, mandibular angle width, and length of the superficial masseteric scar) all show correlated evolution with hypsodonty and the proportion of enamel bands oriented parallel to the chewing stroke. We suggest that changes in the shape of the mandible represent the combined effects of selection for a reorientation of the chewing stroke, so as to emphasize horizontal translation of the teeth, and accommodation of high‐crowned teeth. Our analyses show that the ruminant feeding apparatus is an evolutionary mosaic with its various components showing both correlated and independent evolution. J. Morphol. 275:1093–1102, 2014. © 2014 Wiley Periodicals, Inc.     

Mastication and enamel microstructure in <Emphasis Type="Italic">Cambaytherium</Emphasis>, a perissodactyl-like ungulate from the early Eocene of India

The dentition of Cambaytherium was investigated in terms of dental wear, tooth replacement and enamel microstructure. The postcanine tooth row shows a significant wear gradient, with flattened premolars and anterior molars at a time when the last molars are only little worn. This wear gradient, which is more intensive in Cambaytherium thewissi than in Cambaytherium gracilis, and the resulting flattened occlusal surfaces, may indicate a preference for a durophagous diet. The tooth replacement (known only in C. thewissi) shows an early eruption of the permanent premolars. They are in function before the third molars are fully erupted. During the dominant phase I of the chewing cycle the jaw movement is very steep, almost orthal, with a slight mesiolingual direction and changes into a horizontal movement during phase II. The enamel microstructure shows Hunter-Schreger-bands (HSB) in the inner zone of the enamel. In some teeth the transverse orientation of the HSB is modified into a zig-zag pattern, possibly an additional indicator of a durophagous diet.     

Aspects of masticatory form and function in common tree shrews,Tupaia glis

Tree shrews have relatively primitive tribosphenic molars that are apparently similar to those of basal eutherians; thus, these animals have been used as a model to describe mastication in early mammals. In this study the gross morphology of the bony skull, joints, dentition, and muscles of mastication are related to potential jaw movements and cuspal relationships. Potential for complex mandibular movements is indicated by a mobile mandibular symphysis, shallow mandibular fossa that is large compared to its resident condyle, and relatively loose temporomandibular joint ligaments. Abrasive tooth wear is noticeable, and is most marked at the first molars and buccal aspects of the upper cheek teeth distal to P². Muscle morphology is basically similar to that previously described for Tupaia minor and Ptilocercus lowii. However, in T. glis, an intraorbital part of deep temporalis has the potential for inducing lingual translation of its dentary, and the large medial pterygoid has extended its origin anteriorly to the floor of the orbit, which would enhance protrusion. The importance of the tongue and hyoid muscles during mastication is suggested by broadly expanded anterior bellies of digastrics, which may assist mylohyoids in tensing the floor of the mouth during forceful tongue actions, and by preliminary electromyography, which suggests that masticatory muscles alone cannot fully account for jaw movements in this species.     

Growth allometry of craniomandibular muscles, tendons, and bones in the laboratory rat (Rattus norvegicus): relationships to oromotor maturation and biomechanics of feeding

This study addressed the problem of how growth of craniomandibular muscles, tendons, and bones influences the acquisition of oromotor skills and biomechanics of feeding in the laboratory rat (Rattus norvegicus). Rats representing a 6.6-fold size range were dissected, and muscles, tendons, and mandibles were weighed. Cross-sectional areas of tendons and bones providing attachment surfaces for muscles were estimated. Ontogenetic scaling of craniomandibular muscles, tendons, and bones was described by using linear regression models, and departures from size-required compensations were used to characterize changes in oromotor function. A two-dimensional model was developed which permitted calculation of mechanical advantages of four masticatory muscles; the model was used to show how mandibular growth and tooth eruption influence the biomechanics of rat feeding. Relative to mandible weight, most jaw muscles scaled either isometrically or positively, tendon cross-sectional areas scaled isometrically or negatively, and bone surfaces scaled negatively. With the exception of the superficial masseter and internal pterygoid muscles, mechanical advantages did not change significantly during mandible growth. Growth patterns of craniomandibular muscles, tendons, and bones contribute significantly to changes in morphology and oromotor function.     

Feeding mechanism of Epibulus insidiator (Labridae; Teleostei): Evolution of a novel functional system

The feeding mechanism of Epibulus insidiator is unique among fishes, exhibiting the highest degree of jaw protrusion ever described (65% of head length). The functional morphology of the jaw mechanism in Epibulus is analyzed as a case study in the evolution of novel functional systems. The feeding mechanism appears to be driven by unspecialized muscle activity patterns and input forces, that combine with drastically changed bone and ligament morphology to produce extreme jaw protrusion. The primary derived osteological features are the form of the quadrate, interopercle, and elongate premaxilla and lower jaw. Epibulus has a unique vomero-interopercular ligament and enlarged interoperculo-mandibular and premaxilla-maxilla ligaments. The structures of the opercle, maxilla, and much of the neurocranium retain a primitive labrid condition. Many cranial muscles in Epibulus also retain a primitive structural condition, including the levator operculi, expaxialis, sternohyoideus, and adductor mandibulae. The generalized perciform suction feeding pattern of simultaneous peak cranial elevation, gape, and jaw protrusion followed by hyoid depression is retained in Epibulus. Electromyography and high-speed cinematography indicate that patterns of muscle activity during feeding and the kinematic movements of opercular rotation and cranial elevation produce a primitive pattern of force and motion input. Extreme jaw protrusion is produced from this primitive input pattern by several derived kinematic patterns of modified bones and ligaments. The interopercle, quadrate, and maxilla rotate through angles of about 100 degrees, pushing the lower jaw into a protruded position. Analysis of primitive and derived characters at multiple levels of structural and functional organization allows conclusions about the level of design at which change has occurred to produce functional novelties.     

Enamel ultrastructure and masticatory function in molars of the American opossum, Didelphis virginiana

Maxillary and mandibular molars of the American opossum, Didelphis virginiana L., were viewed in the scanning electron microscope (SEM) after acid-etching or after cutting and acid-etching. Observations were made on enamel prism patterns as they relate to functional properties of the tooth at a particular site. Molars at different stages of wear were also observed under a dissecting microscope; worn surfaces were correlated with function and enamel ultrastructure. Pounding surfaces of molar cusps wear more rapidly than near-vertical shearing surfaces or crushing basins (i.e. the trigon and talonid basin). Pounding surfaces are subjected to abrasion by food and arc not normally involved in tooth-tooth contact. Near-vertical shearing surfaces and basins used for crushing do experience tooth-tooth contact, but are surprisingly more resistant to wear. Prisms at pounding sites approach the occlusal surface at a near 90° angle and are surrounded with very thick interprismatic (IP) enamel parallel to the occlusal surface of the tooth. The pounding pattern is present at tips of cusps and at occlusal surfaces of ridges of the tooth. At near-vertical shearing surfaces, the prisms approach the outer surface obliquely and are surrounded with IP crystals which are perpendicular to the vertical surface. The angle between prismatic and IP enamel in these patterns is 60–90° in a cervical to occlusal direction. In basins of the tooth used principally for crushing and some shearing, IP enamel is perpendicular to the changing slope of the basin and the prisms are usually at a 55–65° angle to the IP enamel. When the pounding and shearing-crushing patterns meet at a ridge, a distinct seam is observed. Pounding forces occur parallel to the long axis of the prisms and perpendicular to the thick IP enamel (i.e. perpendicular to the long axis of the IP crystals) lying on either side of the prisms. Shearing and crushing forces occur at an oblique angle to the prism, and interprismatic enamel is more evenly distributed about the prism. A spiral pattern is found at the bottoms of the trigon and talonid basins, but not at the bottom of the trigonid which is a non-occluding basin. It is concluded that the differential rates of wear of the enamel surfaces are necessary in maintaining the sharp cutting edges and effective crushing basins of the tribosphenic molar, and the ultrastructural arrangements of the enamel prisms are of functional significance.     

Development of the helicoidal plane

In this study of thebelicoidal occlusal plane the relationships between tooth wear, the transverse slopes of mandibular molars and dental arch breadths were examined in 74 pre-contemporary Australian Aboriginal skulls. With increasing age and tooth wear the orientation of the mandibular occlusal surfaces increased towards the buccal. The differential occlusal orientation from first to third molars, present at eruption, tended to increase progressively with tooth wear. These functionally induced changes, together with regional differences in relative breadths of the maxillary and mandibular dental arches, are important in the development of abelicoidal occlusal plane.     

Loss of molar occlusion and mandibular morphology in adults in an ancient human population consuming a coarse diet

The purpose of the study is to investigate the link between number of molar teeth retained in occlusion and mandibular morphology in adults in an ancient, high dental wear human population. The study material comprises skeletons from Mediaeval Wharram Percy, England (N = 50 female, 69 male adults). It was hypothesized that adults retaining fewer occluding molars would show reduction in mandibular dimensions, particularly in the ascending ramus and gonial regions where the main muscles of mastication have their insertions. Molar occlusal status is assessed using the concept of functional units. Mandibular morphology is assessed using a suite of ten linear measurements plus the mandibular angle. Results show no evidence for any association between number of molars retained in occlusion and mandibular angle. There was an association between mandibular size and number of molars retained in occlusion, with smaller mandibular dimensions in those retaining fewer occluding molars. Some measurements were affected more than others so that there was also some shape alteration. Alteration of mandibular dimensions was more clearly demonstrable in females than in males. Only in females could significant reduction in the ascending ramus and gonial regions be demonstrated. Reasons for the apparent difference in response to loss of molar occlusion between male and female mandibles are unclear, but sex differences in bony metabolism mediated by hormonal factors may be implicated. Results suggest that care should be exercised when including mandibles from individuals showing loss of molar occlusion in morphological studies. Am J Phys Anthropol 152:383–392, 2013. © 2013 Wiley Periodicals, Inc.     

Modeling the jaw mechanism of Pleuronichthys verticalis: The morphological basis of asymmetrical jaw movements in a flatfish

Several flatfish species exhibit the unusual feature of bilateral asymmetry in prey capture kinematics. One species, Pleuronichthys verticalis, produces lateral flexion of the jaws during prey capture. This raises two questions: 1) How are asymmetrical movements generated, and 2) How could this unusual jaw mechanism have evolved? In this study, specimens were dissected to determine which cephalic structures might produce asymmetrical jaw movements, hypotheses were formulated about the specific function of these structures, physical models were built to test these hypotheses, and models were compared with prey capture kinematics to assess their accuracy. The results suggest that when the neurocranium rotates dorsally the premaxillae slide off the smooth, rounded surface of the vomer (which is angled toward the blind, or eyeless, side) and are “launched” anteriorly and laterally. The bilaterally asymmetrical trajectory of the upper jaw is determined by the orientation of the “launch pad,” the vomer. During lower jaw depression, the mandibles rotate about their articulations with the quadrate bones of the suspensoria. The quadrato‐mandibular joint is positioned farther anteriorly on the eye side than on the blind side, and this asymmetry deflects the lower jaw toward the blind side. Asymmetry in the articular surfaces of the lower jaw augments this effect. Thus, it appears that fish with intermediate forms of this asymmetrical movement could have evolved from symmetrical ancestors via a few key morphological changes. In addition, similar morphological modifications have been observed in other fish taxa that also produce jaw flexion during feeding, which suggests that there may be convergence in the basic mechanism of asymmetry. J. Morphol. 256:1–12, 2003. © 2003 Wiley‐Liss, Inc.     

Sauropod teeth from the Upper Cretaceous Bissekty Formation of Uzbekistan

The isolated adult teeth of titanosaurian sauropods from the Upper Cretaceous Bissekty Formation at Dzharakuduk, Uzbekistan, differ little in overall structure but show considerable variation in enamel sculpturing and wear patterns. The crown shape of unworn juvenile teeth ranges from lanceolate to conical. Most specimens have enamel texture resembling crumpled paper or completely smooth enamel. Longitudinal grooves along the mesial and distal edges are present on only a few tooth crowns and might be developed on both the labial and lingual sides. Among 252 worn tooth crowns there are eight variants of wear patterns, all possible combinations of 0–2 apical and 0–2 lateral wear facets. The most common is wear pattern A1L0 (one apical facet, no lateral facets; 62.7%). The next most common variant has two apical and no lateral facets (A2L0, 12.3%). These apical wear facets include the primary wear facets, which are produced by an opposing functional tooth, and secondary wear facets, which are produced by a replacing upper tooth coming into contact with the functional lower tooth at a late wear stage. The relative abundance of tooth crowns with two apical wear facets possibly suggests incipient development of a tooth battery in the Bissekty titanosaur.     

Eating with a saw for a jaw: Functional morphology of the jaws and tooth‐whorl in Helicoprion davisii

The recent reexamination of a tooth‐whorl fossil of Helicoprion containing intact jaws shows that the symphyseal tooth‐whorl occupies the entire length of Meckel's cartilage. Here, we use the morphology of the jaws and tooth‐whorl to reconstruct the jaw musculature and develop a biomechanical model of the feeding mechanism in these early Permian predators. The jaw muscles may have generated large bite‐forces; however, the mechanics of the jaws and whorl suggest that Helicoprion was better equipped for feeding on soft‐bodied prey. Hard shelled prey would tend to slip anteriorly from the closing jaws due to the curvature of the tooth‐whorl, lack of cuspate teeth on the palatoquadrate (PQ), and resistance of the prey. When feeding on soft‐bodied prey, deformation of the prey traps prey tissue between the two halves of the PQ and the whorl. The curvature of the tooth‐whorl and position of the exposed teeth relative to the jaw joint results in multiple tooth functions from anterior to posterior tooth that aid in feeding on soft‐bodied prey. Posterior teeth cut and push prey deeper into the oral cavity, while middle teeth pierce and cut, and anterior teeth hook and drag more of the prey into the mouth. Furthermore, the anterior‐posterior edges of the teeth facilitate prey cutting with jaw closure and jaw depression. The paths traveled by each tooth during jaw depression are reminiscent of curved pathways used with slashing weaponry such as swords and knifes. Thus, the jaws and tooth‐whorl may have formed a multifunctional tool for capturing, processing, and transporting prey by cyclic opening and closing of the lower jaw in a sawing fashion. J. Morphol. 276:47–64, 2015. © 2014 Wiley Periodicals, Inc.     

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.