Micro- and macroevolutionary decoupling of cichlid jaws: a test of Liem's key innovation hypothesis

The extent to which elements of functional systems can change independently (modularity) likely influences the diversification of lineages. Major innovations in organismal design, like the pharyngeal jaw in cichlid fishes, may be key to a group's success when they relax constraints on diversification by increasing phenotypic modularity. In cichlid fishes, pharyngeal jaw modifications that enhanced the ability to breakdown prey may have freed their oral jaws from serving their ancestral dual role as a site of both prey capture and prey processing. This functional decoupling that allowed the oral jaws to become devoted solely to prey capture has been hypothesized to have permitted the two sets of cichlid jaws to evolve independently. We tested the hypothesis that oral and pharyngeal jaw mechanics are evolutionarily decoupled both within and among Neotropical Heroine cichlids. In the trophically polymorphic species Herichthys minckleyi, molariforms that exhibit enlarged molarlike pharyngeal jaw teeth were found to have approximately 400% greater lower jaw mass compared to H. minckleyi with the alternative papilliform pharyngeal morphology. However, oral jaw gape, lower jaw velocity ratios, anterior jaw linkage mechanics, and jaw protrusion did not differ between the morphotypes. In 40 other Heroine species, there was a weak correlation between oral jaw mechanics and pharyngeal jaw mass when phylogenetic history was ignored. Yet, after expansion of the cytochrome b phylogeny for Heroines, change in oral jaw mechanics was found to be independent of evolutionary change in pharyngeal jaw mass based on independent contrasts. Evolutionary decoupling of oral and pharyngeal jaw mechanics has likely played a critical role in the unparalleled trophic diversification of cichlid fishes.     

Aspects of the Feeding and Reproductive Biology of the Lake Tanganyikan Cichlid, Lamprologus ornatipinnis (Pisces, Cichlidae)

This project investigated the reproductive and feeding biology of Lamprologus ornatipinnis. Specimens were collected monthly from North Bay, Mbita Island, Zambia for a year. Dentition was examined under a scanning electron microscope, stomach contents were analysed using the Index of Relative Importance (IRI) and mean Gonado-somatic Indices (GSI) were calculated for each month to reveal spawning peaks. Both males and females possess an outer row of six (premaxilla) and eight (dentary) enlarged canines. The remaining inner rows consist of small, recurved canine-like teeth. The lower pharyngeal bone possesses enlarged centrally placed molars with an increasing number of smaller bevelled teeth laterally. This dentition is very similar to a typical benthic arthropod and mollusc eater. Stomach content analysis revealed that L. ornatipinnis feeds predominantly on Chironomidae, Copepoda and Ostracoda. The importance of these prey items in the diet differed significantly between the months sampled. Percentage volume of Copepoda and Cyclopoida was greater in female stomachs than males and the % number of Chironomidae greater in males than females. Gonadosomatic Index (GSI) values indicated males and females breed throughout the year. Females, however have a significant peak (p<0.05) in breeding activity from July to October that coincides with the increased abundance of plankton at this time.     

Terrestrial feeding in the Mudskipper Periophthalmus (Pisces: Teleostei): A cineradiographic analysis

Mudskipping gobies (Periophthalminae) are among the most terrestrial of amphibious fishes. Specializations associated with terrestrial prey capture and deglutition have been studied in Periophthalmus koelreuteri by light and X-ray cinematography which permits direct visualization of pharyngeal jaw movement during deglutition. Anatomical specializations of the pharyngeal jaws are described and include depressible teeth, a large ventral process on ceratobranchial five, and muscular modifications.
Multiple terrestrial feedings occur by Periophthalmus without a return to the water, and cineradiography reveals that the buccal cavity is often filled with air during terrestrial excursions in contrast to some previous hypotheses. Transport of the prey into the oesophagus occurs primarily by anteroposterior movement of the upper pharyngeal jaw. The lower pharyngeal jaw plays a limited role in food transport and may serve primarily to hold and position prey. The bite between upper and lower pharyngeal jaws occurs between the anterior teeth, and both jaws are protracted together during raking of food into the oesophagus. Functional specializations correlated with terrestrial feeding include obligatory use of pharyngeal jaws for swallowing even small prey items and positioning of the prey in the pharynx by pharyngeal jaw and hyoid movements alone.
This analysis of terrestrial feeding allows hypotheses of design constraints imposed by the aquatic medium on fishes to be raised and tested.     

The phylogenetic significance of bone types in euteleost fishes

The paradentary is a small, sometimes dentigerous element in the lower jaw of some atherinomorph neoteleost fishes. Identification of the paradentary as a neomorphic, perichondrally ossified bone prompted re-examination of theories of the association of bone and teeth in teleost fishes. Teeth on a chondral lower jaw bone might be explained simply by epidermal-mesodermal interactions. Since the work of Kolliker in 1859, it has been known that there are two basic types of bone in teleost fishes: cellular bone, characterized by a matrix that has enclosed osteoblasts or osteocytes; and acellular bone, characterized by a relatively featureless matrix that lacks these bone-forming cells. Cellular bone is typical of lower teleosts, whereas acellular bone is typical of higher teleosts. Ontogenetic data indicate that acellular bone is derived relative to cellular bone. Even though identification of cellular and acellular bone can be made readily with histological preparations, acellular bone has been used infrequently as a character in analyses of teleost phylogeny. Acellular bone is considered here to be a derived character within teleost fishes. It is found in all Neoteleostei as well as some, but not all Salmoniformes. Independent of studies of bone, derived types of teeth in teleosts have been described in terms of their failure to become completely mineralized. Acellular bone and teeth of higher teleosts share several properties, including a large fraction of collagen. Teleosts lack a parathyroid gland; bone type is critical to the mechanism of calcium regulation. It is proposed that the character of acellular bone be incorporated into phylogenetic analyses of teleost fishes by correlating it with derived types of tooth structure.     

Scanning microscopy of pharyngeal and oral teeth of the teleost Tilapia mossambica (Peters)

The pharyngeal and oral teeth of the fish Tilapia mossambica (Peters) were examined with a scanning microscope. It appeared that the dorsal pharyngeal teeth form a peculiar hooklike extension at the tip, whereas the ventral pharyngeal teeth tend to curve in a posterior direction. The two lateral flanges at the tip of the ventral teeth are probably the areas of contact with the dorsal teeth when the latter are pressed down during sound production or feeding. However, the oral teeth develop along a different line. A part from villiform teeth the upper and lower jaws also develop tricuspid and bicuspid oral teeth, with the bicuspids concentrated mainly along the outer edge of the jaw.     

Morphological and videofluorographic study of the hyoid apparatus and its function in the rabbit (Oryctolagus cuniculus)

The anatomy of the hyoid apparatus and positional changes of the hyoid bone during mastication and deglutition are described in the New Zealand White rabbit (Oryctolagus cuniculus). A testable model is constructed to predict the range of movement during function of the hyoid, a bone entirely suspended by soft tissue. Frame-by-frame analysis of a videofluorographic tape confirms the accuracy of the prediction through observation of hyoid bone excursion during oral behavior. During chewing, translation of the hyoid bone is diminutive and irregular, lacking a clearly discernible path of excursion. However, some movements of the hyoid occur with regularity. During fast opening, anterodorsal movement of the hyoid is interrupted with an abrupt posteroventral depression when the bolus is moved posteriorly toward the cheek teeth by the tongue. This clockwise rotation (when viewed from the right side) of the hyoid accompanies jaw opening and is reversed (posteroventral movement) for the jaw closing sequence. Lateral movements of the hyoid may be slightly coupled to mandibular rotation in the horizontal plane. The findings suggest that the hyoid bone maintains a relatively static position during the dynamics of chewing. The primary function would be to provide a stable base for the movements of the tongue. Another possible function would be to control the position of the larynx within the pharyngeal cavity. Some characteristic features of the rabbit hyoid apparatus may be consequential to relatively erect posture and a saltatory mode of locomotion.     

An Ancient Gene Network Is Co-opted for Teeth on Old and New Jaws

Vertebrate dentitions originated in the posterior pharynx of jawless fishes more than half a billion years ago. As gnathostomes (jawed vertebrates) evolved, teeth developed on oral jaws and helped to establish the dominance of this lineage on land and in the sea. The advent of oral jaws was facilitated, in part, by absence of hox gene expression in the first, most anterior, pharyngeal arch. Much later in evolutionary time, teleost fishes evolved a novel toothed jaw in the pharynx, the location of the first vertebrate teeth. To examine the evolutionary modularity of dentitions, we asked whether oral and pharyngeal teeth develop using common or independent gene regulatory pathways. First, we showed that tooth number is correlated on oral and pharyngeal jaws across species of cichlid fishes from Lake Malawi (East Africa), suggestive of common regulatory mechanisms for tooth initiation. Surprisingly, we found that cichlid pharyngeal dentitions develop in a region of dense hox gene expression. Thus, regulation of tooth number is conserved, despite distinct developmental environments of oral and pharyngeal jaws; pharyngeal jaws occupy hox-positive, endodermal sites, and oral jaws develop in hox-negative regions with ectodermal cell contributions. Next, we studied the expression of a dental gene network for tooth initiation, most genes of which are similarly deployed across the two disparate jaw sites. This collection of genes includes members of the ectodysplasin pathway, eda and edar, expressed identically during the patterning of oral and pharyngeal teeth. Taken together, these data suggest that pharyngeal teeth of jawless vertebrates utilized an ancient gene network before the origin of oral jaws, oral teeth, and ectodermal appendages. The first vertebrate dentition likely appeared in a hox-positive, endodermal environment and expressed a genetic program including ectodysplasin pathway genes. This ancient regulatory circuit was co-opted and modified for teeth in oral jaws of the first jawed vertebrate, and subsequently deployed as jaws enveloped teeth on novel pharyngeal jaws. Our data highlight an amazing modularity of jaws and teeth as they coevolved during the history of vertebrates. We exploit this diversity to infer a core dental gene network, common to the first tooth and all of its descendants.     

Dentition in the African catfishes Andersonia (Amphiliidae) and Siluranodon (Schilbeidae) previously considered toothless

Based on light and scanning electron microscopic examination of their morphology, the dentition on both the premaxilla and dentary of Andersonia (Amphiliidae) and Siluranodon (Schilbeidae) catfishes is described from samples taken from tributaries of the White Nile in south‐western Ethiopia. These monotypic African genera were previously believed to lack teeth on the lower jaw in Andersonia and on both jaws in Siluranodon . Siluranodon exhibits an ontogenetic reduction: teeth were less frequently found in larger individuals than in smaller ones. In contrast to the adults of all other schilbeids, whose oral teeth are arranged in multiserial (or at least, biserial) bands, Siluranodon has uniserial teeth on both the premaxilla and the dentary. The adaptive, ontogenetic and phylogenetic aspects of jaw‐tooth reduction in catfishes are discussed.     

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.     

Regeneration of ankylosed teeth in the adult frog premaxilla.

The relationship between the ankylosed amphibian tooth and regeneration of a jaw segment was studied. A section of the premaxilla was removed in 95 young leopard frogs. Subsequent would healing was observed at intervals of 0-180 days. The dental lamina formed new teeth and by 90 days, teeth in varying stages of development could be seen extending across the wound segment. Teeth within the wound grew to normal size and shape and were replaced by their successors without support of underlying jaw bone which grew in later and often was incomplete.     

A new genus and species of Sarcoglanidinae from the Río Mamoré, Amazon Basin, with comments on subfamilial phylogeny (Teleostei, Trichomycteridae)

Stenolicmus sarmientoi is described as a new genus and species of the trichomycterid subfamily Sarcoglanidinae, from the RíAo Mamoré Basin, Bolivia. It can be distinguished from all other sarcoglanidines by: five-rayed pectoral fin; elongate body shape (HL about 15% of SL); absence of fontanelles on cranial roof; well-developed patches of opercular and interopercular odontodes (five or six on each bone); numerous accessory caudal-fin rays (13 dorsal and 11 ventral); and presence of extensive dark pigmentation on the surface of the body. Three synapomorphies support a hypothesis that the new taxon forms a monophyletic group with other sarcoglanidines: presence of an anterior ossification of the palatine cartilage; quadrate with a dorsal expansion directed posteriorly; and premaxilla with a long lateral process. Stenolicmus lacks some synapomorphies shared by Stauroglanis, Malacoglanis and Sarcoglanis , and shares some other synapomorphies exclusively with Sarcoglanis and Malacoglanis. These character distributions make it uncertain whether Stenolicmus is the sister group of all other sarcoglanidines or only of Malacoglanis plus Sarcoglanis , because the two positions imply equally parsimonious cladograms.     

Structure of the skin of Agonus cataphractus (Teleostei)

The skin of the scuted teleost Agonus cataphractus has been investigated by histochemical methods, SEM and TEM. The anterior dorsal skin bears tubercles of epidermis overlying tiny ossifications (scutelets) superficial to the main scutes. The epidermis secretes a cuticular layer containing acidic non-sulphated glycoproteins, but there are no mucous goblet cells in the external skin. Non-mucous sacciform cells of two types are present in the epidermis, also numerous chloride cells. Scanning electron microscopy reveals variation in the microridge pattern of superficial epithelial cells, thought to relate to arrival at the surface and secretion of the cuticle. The major scutes overlap anteriorly, contrary to the normal arrangement of scales, indicating that they are secondary ossifications. The type of mineralization is similar to that of acellular bone. The scutes are set directly in the collagen of the dermis. They have a girdered structure with radial and cross bars, inserting on both faces of a thin plate. The interstices are occupied by unmineralized collagen, and extrinsic collagen bundles impinge on the bone. Non-mineralized parts of the dermis contain tracts of microfibrils in addition to collagen; these are best developed in the flexible gular skin and in the barbels and are interpreted as elastic tissue, although an amorphous component was not seen. The barbels have a core of connective tissue without a cartilaginous skeleton and bear taste buds and numerous chloride cells.     

Anatomy of a hemiramphid pharyngeal mill with reference to Arrhamphus sclerolepis krefftii (Steindachner) (Teleostei: Hemiramphidae)

The structure and function of the pharyngeal jaw apparatus (PJA) and postpharyngeal alimentary tract of Arrhamphus sclerolepis krefftii, an herbivorous hemiramphid, were investigated by dissection, light and scanning electron microscopy, and X-ray analysis of live specimens. A simple model of PJA operation is proposed, consisting of an adductive power stroke of the third pharyngobranchial that draws it posteriorly while the fifth ceratobranchial is adducted, and a return stroke in which the third pharyngobranchial bone is drawn anteriorly during abduction of the fifth ceratobranchial. Teeth in the posteromedial region of the PJA are eroded into an occlusion zone where the teeth of the third pharyngobranchial are spatulate incisiform and face posteriorly in opposition to the rostrally oriented spatulate incisiform teeth in the wear zone of the fifth ceratobranchial. The shape of the teeth and their pedestals (bone of attachment) is consistent with the model and with the forces likely to operate on the elements of the PJA during mastication. The role of pharyngeal tooth replacement in maintaining the occlusal surfaces in the PJA during growth is described. The postpharyngeal alimentary tract of A. sclerolepis krefftii comprises a stomachless cylinder that attenuates gradually as it passes straight to the anus, interrupted only by a rectal valve. The ratio of gut length to standard length is about 0.5. Despite superficial similarities to the cichlid PJA (Stiassny and Jensen [1987] Bull Mus Comp Zool 151:269-319), the hemiramphid PJA differs in the fusion of the third pharyngobranchial bones, teeth in the second pharyngobranchials and the fifth ceratobranchial face anteriorly, the presence of a slide-like diarthroses between the heads of the fourth epibranchials and the third pharyngobranchial, the occlusion zone of constantly wearing teeth, and the unusual form of the muscularis craniopharyngobranchialis. The functional relationship between these structures is explained and the consequence for the fish of a complex PJA and a simple gut 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.     

Fabrosauridae,the basal family of ornithischian dinosaurs (Reptilia: Ornithopoda)

In fabrosaurids the upper jaw is flat and the lower jaw is slender so the ’cheek’ teeth are marginal and not inset as is the case in all other ornithischian dinosaurs. The ’cheek’ teeth of fabrosaurids have anteroposteriorly expanded crowns but lack wear surfaces formed by tooth to tooth contact. Two genera are recognized from the Triassic-Jurassic boundary of Lesotho with good material previously referred toFabrosaurus as a new genus that represents the most conservative ornithopod described to date. The anatomy ofNanosaurus (Upper Jurassic, U.S.A.) andEchinodon (Jurassic-Cretaceous boundary, England) is redescribed; in both genera the tooth bearing bone of the lower jaw is deepened posteriorly and inEchinodon there is a true canine tooth in the upper jaw.     

Variation in body size and trophic morphology within and among genetically differentiated populations of the cichlid fish, Metriaclima zebra, from Lake Malawi

1. The cichlid fish Metriaclima zebra , common in Lake Malawi, feeds by filtering plankton from the water and by brushing items from sediment covered substrata. It inhabits isolated rocky reefs among which community structure, resource availability and gene pools are likely to differ. We speculated that body size and trophic morphology of M. zebra might vary concomitantly.
2. We quantified the extent of genetic, body size and trophic variation within and between populations of M. zebra from southern Lake Malawi. Specifically, we tested the hypotheses that: (i) local populations are genetically differentiated, (ii) local populations differ in jaw morphology, dentition and standard length (SL), and (iii) variation in size is correlated with variation in trophic morphology.
3. Local populations of M. zebra differed in mean SL and were genetically differentiated. Moreover, populations exhibited dissimilar oral jaw morphologies and dentitions, perhaps related to differences in feeding biology. Variation in jaw shape was largely restricted to the curvature of the distal tip of the dentary. Populations were characterised by individuals with oblique, upward or downward directed gapes. Dental patterns differed in the proportion of unicuspid teeth in all rows of each jaw (dentaries and premaxillae) and the spacing of teeth in affected rows.
4. Within populations, jaw and tooth shapes were correlated with body size. Smaller individuals possessed upward curving jaws and closely packed multicusped teeth, while larger individuals exhibited relatively downward-directed jaws with increasing numbers of widely spaced unicuspid teeth.
5. Metriaclima zebra populations have increased in mean SL over the last decade, in contrast to a decline among Lake Malawi pelagic cichlids. Differences in size may contribute to variation in trophic morphology and may track local environmental dynamics in this lacustrine system.     

Inferring parrotfish (Teleostei: Scaridae) pharyngeal mill function from dental morphology, wear, and microstructure

Morphology, occlusal surface topography, macrowear, and microwear features of parrotfish pharyngeal teeth were investigated to relate microstructural characteristics to the function of the pharyngeal mill using scanning electron microscopy of whole and sectioned pharyngeal jaws and teeth. Pharyngeal tooth migration is anterior in the lower jaw (fifth ceratobranchial) and posterior in the upper jaw (paired third pharyngobranchials), making the interaction of occlusal surfaces and wear-generating forces complex. The extent of wear can be used to define three regions through which teeth migrate: a region containing newly erupted teeth showing little or no wear; a midregion in which the apical enameloid is swiftly worn; and a region containing teeth with only basal enameloid remaining, which shows low to moderate wear. The shape of the occlusal surface alters as the teeth progress along the pharyngeal jaw, generating conditions that appear suited to the reduction of coral particles. It is likely that the interaction between these particles and algal cells during the process of the rendering of the former is responsible for the rupture of the latter, with the consequent liberation of cell contents from which parrotfish obtain their nutrients.     

Shape analysis of the jaws between two minnow species over ontogeny

This study compares sand shiner (Notropis stramineus ) and silverjaw (Ericymba buccata ) minnows, in terms of the morphological shape changes of the upper, lower, and pharyngeal jaws over ontogeny. These two species of minnows initially feed on midge larvae and undergo an ontogenic prey shift. The traditional morphometrics measured—total length, snout‐to‐vent length, eye diameter, premaxilla length, lower jaw length, gape—were regressed onto total length to test for allometry. Digital pictures were processed with tpsDig and further analyzed with MorphoJ utilizing a regular geometric morphometrics procedure using principle component analyses. We examined gut contents for 16 fish of each species. For the silverjaw minnows, we found all jaw variables to exhibit positive allometric growth with increasing total length, while most of the jaw variables for the sand shiner exhibited negative allometric growth with increasing total length. This correlates with an ontogenic prey shift for both species. Sand shiner minnows have been found to be more omnivorous, feeding on algae later in life, while silverjaw minnows undergo a prey shift to larger invertebrates. These species lack oral dentition causing an increased reliance on the pharyngeal apparatus. Principle component analyses revealed elongation of pharyngeal jaw elements in the silverjaw minnows and a relative shortening and bulking of pharyngeal jaws in the sand shiner minnows. The ontogenic dietary shifts observed in these two species provide possible explanation for the morphological changes over ontogeny in jaw elements, which are likely enabling these species to occupy the same habitat with little niche overlap.     

Structure and formation of ankylosis in Xenopus laevis

The structure of ankylotic teeth in Xenopus laevis was studied by light, transmission, and scanning electron microscopy as well as by microradiography in decalcified and undecalcified specimens. The mature teeth of Xenopus laevis are calcified from the crown to the base, fused to the jaw bone, and have no uncalcified area, such as a fibrous ring separating the tooth into the crown and pedicle. Microradiography shows that the mature tooth and jaw bone appear as an X-ray opaque area, except for the basal region of the dentine. This region is composed of an X-ray translucent area and an X-ray opaque thin layer on the lingual side of the translucent area. The mature tooth is composed of two differently calcified areas: (1) a highly calcified area, which makes up almost all of the tooth and contains a thin layer of the basal dentine on the lingual side, and (2) a lowly calcified basal dentine, which is fused to the jaw bone. Therefore, the lowly calcified area does not completely separate the dentine and jaw bone. Repeating banding patterns among the collagen fibrils differ among the dentine-forming area and the matrices of dentine and jaw bone. During the formation of ankylosis of the tooth germ, collagen bundles in the dentine-forming area accumulate directly on the surface of the jaw bone. Consequently, the mature teeth of Xenopus laevis fuse to the jaw bone directly without the mediation of the other structures.     

Surface changes in the naturally ankylosed teeth of the frog Rana pipiens during growth and maturation: an SEM study

An SEM study of the surface morphology of the major stages of mature and developing teeth of the leopard frog was made using anorganic preparations of the teeth and jaws. After initial development, the crown area changed little during subsequent tooth eruption, ankylosis and maturation. The thin enamel covering extended further down the shaft than expected. After ankylosis, the surfaces of the tooth continued to mature. The unmineralized gap between the crown and the pedestal, which is prominent in most amphibians, gradually filled in as the ankylosed tooth aged. The upper portion of the pedestal initially formed a dentine surface which was globular in appearance due to partial calcification of the surface collagen fibres but became smooth with uniformly calcified fibres as the ankylosed tooth matured. The lower portion of the pedestal was more variable and there was a gradual transition of dentine into a more cellular, bone-like tissue which contained lacunae and larger fibre bundles. This bone-like tissue was very distinct in surface morphology from the bone of the adjacent jaw, and as the tooth matured it changed from a coarse, woven appearance to one more like lamellar bone. Resorption bays were present in both the dentine and bony areas of teeth which were being shed. During development, the pedestal, which attaches the tooth to the jaw, formed as a separate calcification site and did not form a complete ring until fusion of its buccal surface with that of the overlying crown. A bony buccal lip formed early as part of the pedestal.