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.     

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.     

Viviparity in the halfbeak genera Dermogenys and Nomorhamphus (Teleostei: Hemiramphidae)

Gravid ovaries were examined histologically from two species of Nomorhamphus and 21 populations of Dermogenys. In addition, changes in dry-weight throughout gestation are provided for 15 populations. The ovaries are paired organs running along the lateral body wall and are separated along most of their length. In all specimens examined, embryos are fertilized within the ovarian follicle. Viviparity in these species is divided herein into five categories designated types I–V. In types I and II the entire gestation period is intrafollicular, whereas in types III–V only the early stages of gestation are intrafollicular with the major period of development occurring in the ovarian lumen (intraluminal). Type I is characterized by the retention of a large amount of yolk throughout gestation. Superfetation is not observed. Populations of D. pusilla from Vietnam and Thailand decrease in dry-weight throughout gestation. This, coupled with the slight vascularization of the yolk sac, suggests strict lecithotrophy. Populations of D. pusilla from Singapore and Bangladesh undergo an increase in dry weight and exhibit an increased vascularization of the yolk sac, suggesting a form of unspecialized matrotrophy. Type II is characterized by a small amount of yolk, an expansion of the coelomic cavity and pericardial sac, and a simple cuboidal epithelium on the general body surfaces. Superfetation occurs with up to three broods present within a single ovary. Dermogenys pusilla from Sabah, D. orientalis and Dermogenys sp. (Sulawesi) exhibit the type II form of viviparity. Dermogenys vivipara from the eastern Philippine islands of Culion and Busuanga exhibit characteristics considered intermediate between type I and II. These results are compared with those from other viviparous species exhibiting intrafollicular gestation. In species with types III–V (intraluminal gestation), developing oocytes are restricted to a distinct ridge of ovigerous tissue extending along the entire length of the ovary. Two species, D. viviparus (Luzon, Philippines) and Dermogenys sp. (Luzon) have the type III form of viviparity. In this form, oocytes are small (0.8–1.0 mm) with little yolk reserves and embryos, covered with a simple cuboidal epithelium and possessing an expanded belly sac, are retained within the follicles until a late fin-bud stage. Type III embryos found within the ovarian lumen have a greatly expanded belly sac and remain covered by a simple cuboidal epithelium until parturition. Superfetation is present in these species with two broods observed simultaneously within a single ovary. Five species, D. megarrhamphus, D. weberi, D. viviparus (Jolo, Philippines), Nomorhamphus sp. (Sulawesi), and N. towoetii, were observed with the type IV form of viviparity. Embryos in this category are evacuated into the ovarian lumen prior to a fin-bud stage and retain a large yolk mass throughout development. Superfetation is absent in these species. A differentform of viviparity (type V) is present in D. ebrardtii in which embryos appear to obtain nutrients through a form of oophagy and aldelphophagy (feeding on developing oocytes or less-developed siblings). In all specimens with intraluminal development, atretic oocytes within the ovigerous ridge are abundant. These findings support the hypothesis that current species and generic limits may be artificial and underscores the potential of histological evidence for phylogenetic analysis of this group. J. Morphol. 234:295–317, 1997. © 1997 Wiley-Liss, Inc.     

Unique dentition of rhynchosaurs and their two-phase success as herbivores in the Triassic

Rhynchosaurs were key herbivores over much of the world in the Middle and Late Triassic, often dominating their faunas ecologically, and much of their success may relate to their dentition. They show the unique ankylothecodont mode of tooth implantation, with deep roots embedded in the bone of the jaw and low crowns that were rapidly worn down in use. During growth, the main area of oral food processing, located in the middle and posterior portions of the occlusal surfaces of the jaws, moved posteriorly relative to the anterior tips of the jaws, which curved up. As the maxilla and dentary grew by addition of new bone posteriorly, the dental lamina fed in new teeth at the back of the tooth rows. CT scanning of the holotype skull of Bentonyx sidensis from the Middle Triassic of England reveals previously concealed details of the dentition. Together with new dentary material from the same location, this has enabled us to examine the tooth replacement process and elucidate ontogenetic changes in dentition and jaw morphology as the animals aged. There were major changes in rhynchosaur anatomy and function through their evolutionary history, with the early forms of the Middle Triassic dying out before or during the Carnian Pluvial Episode (233–232 Ma), and the subclade Hyperodapedontinae, with broad skulls and adaptations to chop tough vegetation, subsequently diversifying worldwide in a successful ecological expansion until their global extinction 227–225 Ma.     

Phylogenetic systematics of the viviparous halfbeak genera Dermogenys and Nomorhamphus (Teleostei:Hemiramphidae:Zenarchopterinae)

Among the 13 genera and over 100 species of halfbeaks, three genera - Dermogenys, Nomorhamphus and Hemirhamphodon – are internally fertilized and viviparous. These genera belong to a more inclusive clade, the Zenarchopterinae, that also includes Zenarchopterus , inferred to be internally fertilized and to lay fertilized eggs, and the monotypic Tondanichthys , also inferred to be internally fertilized. Whereas the Hemiramphidae are distributed worldwide, internally fertilized halfbeaks are restricted to Southeast Asia. Recent data from histological surveys of the gonads of both males and females as well as embryonic modifications associated with viviparity have been combined here with osteological characters in a phylogenetic analysis. Results indicate overwhelming support for a sister-group relationship between Hemirhamphodon and {Dermogenys + Nomorhamphus). Monophyly of the Dermogenys + Nomorhamphus clade is also well supported. These results confirm earlier suggestions that Dermogenys , as previously defined, is paraphyletic. Within the Dermogenys + Nomorhamphus clade, two monophyletic clades are supported:one comprises ten species including four new species (Dermogenys bruneiensis, Dermogenys robertsi, Dermogenys palawanensis and Dermogenys collettei) and the other comprises 13 species including three undescribed species (Nomorhamphus rossi, Nomorhamphus pinnimaculata and Nomorhamphus manifesta). Diagnoses for the species of Dermogenys and Nomorhamphus , as well as a natural classification for the included species, are presented.     

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.     

Autoradiographic confirmation of mucus-mediated absorption in the halfbeak Hyporhamphus regularis ardelio (Whitley, 1931)

In an investigation of the relationship between acidic glycoproteins released in the pharynx and oesophagus of Hyporhamphus regularis ardelio and 'juice' released from Zostera capricorni by pharyngeal milling, autoradiographs of thin sections of guts of fish fed C¹⁴ labelled seagrass leaves revealed high emission densities in the acidic glycoprotein sheath surrounding the ingesta. This indicates that 'juice' liberated from seagrass is taken up by the mucus.     

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.     

Development of the pharyngeal arch skeleton in Catostomus commersonii (Teleostei: Cypriniformes)

Skeletal elements of the gill arches of adult cypriniform fishes vary widely in number, size, and shape and are important characters in morphologically based phylogenetic studies. Understanding the developmental basis for this variation is thus phylogenetically significant but also important in relation to the many developmental genetic and molecularly based studies of the early developing and hence experimentally tractable gill arches in the zebrafish, a cyprinid cypriniform. We describe the sequence of the chondrification and ossification of the pharyngeal arches and associated dermal bones from Catostomus commersonii (Catostomidae, Cypriniformes) and make selected comparisons to other similarly described pharyngeal arches. We noted shared spatial trends in arch development including the formation of ventral cartilages before dorsal and anterior cartilages before posterior. Qualitatively variable gill arch elements in Cypriniformes including pharyngobranchial 1, pharyngobranchial 4, and the sublingual are the last such elements to chondrify in C. commersonii. We show that the sublingual bone in C. commersonii has two cartilaginous precursors that fuse and ossify to form the single bone in adults. This indicates homology of the sublingual in catostomids to the two sublingual bones in the adults of cobitids and balitorids. Intriguing patterns of fusion and segmentation of the cartilages in the pharyngeal arches were discovered. These include the individuation of the basihyal and anterior copula through segmentation of a single cartilage rod, fusion of cartilaginous basibranchials 4 and 5, and fusion of hypobranchial 4 with ceratobranchial 4. Such “fluidity” in cartilage patterning may be widespread in fishes and requires further comparative developmental studies. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

The shark fauna from the Middle Triassic (Anisian) of North-Western Nevada

The shark fauna from the Anisian of Nevada is dominated by durophagous hybodontiforms but also shows an important neoselachian component. Two new species of hybodontiform sharks, Acrodus cuneocostatus and Polyacrodus bucheri , are described in addition to a new neoselachian taxon: Mucrovenator minimus. The enameloid of the teeth of Acrodus and Polyacrodus comprises two layers, an outer compact layer and an inner bundled layer. For the typical three-layered enameloid of neoselachian teeth, we propose to replace the terms parallel-fibred enameloid and tangled-fibred enameloid by the more appropriate parallel-bundled and tangled-bundled enameloid.     

Diversity and convergences in the evolution of feeding adaptations in ankylosaurs (Dinosauria: Ornithischia)*

Ankylosaurian dinosaurs were low-browsing quadrupeds that were traditionally thought of as simple orthal pulpers exhibiting minimal tooth occlusion during feeding, as in many extant lizards. Recent studies, however, have demonstrated that effective chewing with tooth occlusion and palinal jaw movement was present in some members of this group. Qualitative and quantitative analysis of feeding characters (i.e. craniodental features, tooth wear patterns, origin and insertion of jaw adductors) reveal at least three different jaw mechanisms during the evolution of Ankylosauria. Whereas, in basal members, food processing was restricted to simple orthal pulping, in late Early and Late Cretaceous North American and European forms a precise tooth occlusion evolved convergently in many lineages (including nodosaurids and ankylosaurids) complemented by palinal power stroke. In contrast, Asian forms retained the primitive mode of feeding without any biphasal chewing, a phenomenon that might relate to the different types of vegetation consumed by these low-level feeders in different habitats on different landmasses. Further, a progressive widening of the muzzle is demonstrated both in Late Cretaceous North American and Asian ankylosaurs, and the width and general shape of the muzzle probably correlates with foraging time and food type, as in herbivorous mammals.     

Morphological analysis of the pharyngeal jaws in two populations of Lebias fasciata Valenciennes, 1821 (Teleostei: Cyprinodontidae)

The study of the pharyngeal jaws in two geographically isolated Italian populations of Lebias fasciata indicated the presence of two phenotypes: the Adriatic phenotype with a large ceratobranchial V and upper and lower pharyngeal jaws bearing few large teeth and the Sicilian phenotype with a smaller ceratobranchial V and pharyngeal jaws with smaller and more numerous teeth. The morphological variations of pharyngeal jaws should be interpreted as a result of the geographical isolation of these two populations. J. Morphol. 241:107–114, 1999. © 1999 Wiley‐Liss, Inc.     

Four rows of pharyngeal teeth in an aberrant specimen of the small African barb Barbus paludinosus(Cyprinidae): novelty or atavistic alteration?

An extra fourth row of pharyngeal teeth was found on both pharyngeal arches in a specmen of the small African barb, Barbus paludinosus , a species normally exhibiting three rows of pharyngeal teeth. Possible phylogenetic significance of this alteration is discussed in connection with occurrence of extra fourth rows of pharyngeal teeth in other cyprinid lineages.     

Taxonomy,morphology, masticatory function and phylogeny of heterodontosaurid dinosaurs

Heterodontosaurids comprise an important early radiation of small-bodied herbivores that persisted for approximately 100 My from Late Triassic to Early Cretaceous time. Review of available fossils unequivocally establishes Echinodon as a very small-bodied, late-surviving northern heterodontosaurid similar to the other northern genera Fruitadens and Tianyulong. Tianyulong from northern China has unusual skeletal proportions, including a relatively large skull, short forelimb, and long manual digit II. The southern African heterodontosaurid genus Lycorhinus is established as valid, and a new taxon from the same formation is named Pegomastax africanus gen. n., sp. n. Tooth replacement and tooth-to-tooth wear is more common than previously thought among heterodontosaurids, and in Heterodontosaurus the angle of tooth-to-tooth shear is shown to increase markedly during maturation. Long-axis rotation of the lower jaw during occlusion is identified here as the most likely functional mechanism underlying marked tooth wear in mature specimens of Heterodontosaurus. Extensive tooth wear and other evidence suggests that all heterodontosaurids were predominantly or exclusively herbivores. Basal genera such as Echinodon, Fruitadens and Tianyulong with primitive, subtriangular crowns currently are known only from northern landmasses. All other genera except the enigmatic Pisanosaurus have deeper crown proportions and currently are known only from southern landmasses.     

Estimating age from adult occlusal wear: A modification of the miles method

The Miles method of age estimation relies on molar wear to estimate age and is widely used in bioarcheological contexts. However, because the method requires physical seriation and a sample of subadults to estimate wear rates it cannot be applied to many samples. Here, we modify the Miles method by scoring occlusal wear and estimating molar wear rates from adult wear gradients in 311 hunter‐gatherers and provide formulae to estimate the error associated with each age estimate. A check of the modified method in a subsample (n = 22) shows that interval estimates overlap in all but one case with age categories estimated from traditional methods; this suggests that the modifications have not hampered the ability of the Miles method to estimate age even in heterogeneous samples. As expected, the error increases with age and in populations with smaller sample sizes. These modifications allow the Miles method to be applied to skeletal samples of adult crania that were previously only amenable to cranial suture age estimation, and importantly, provide a measure of uncertainty for each age estimate. Am J Phys Anthropol 149:181–192, 2012. © Wiley Periodicals, Inc.     

Rethinking incisor size and diet in anthropoids: diet, incisor wear and incisor breadth in the African apes

In a seminal study Hylander (1975) concluded that the length of the incisor row in catarrhines considered frugivores is longer relative to body mass than in those classified as folivores. Assuming that large fruits require greater incisal processing than do leaves, stems, berries, and seeds, he argued that the larger incisors of frugivores increased their resistance to wear. The present analysis examines diet, incisor wear, and incisor crown breadth in cranial samples of western lowland gorillas and chimpanzees. Incisor wear rate was assessed on the basis of the extent of incisor crown reduction observed at sequential stages of first molar wear. Incisor metrics were obtained from the unworn teeth of juveniles. Results suggest that incisor wear is greater in the more folivorous western lowland gorillas than in more frugivorous chimpanzees. Moreover, incisor crown dimensions do not differ appreciably among African apes. These findings fail to support the hypothesis that slower wear rates are associated with broader incisor crowns, and raise new questions regarding the significance of incisor row length in anthropoids.     

Ontogeny, anatomy and attachment of the dentition in mosasaurs (Mosasauridae: Squamata)

Aspects of mosasaur dental ontogeny are well preserved in many fossils of these giant marine squamates. Replacement teeth on the tooth-bearing elements (TBEs) first appear as small enamel crowns positioned posterolingual to the attached tooth (posterolabial for the pterygoid). Several developing crowns, of progressively larger size, are aligned in rows relating to a specific tooth position. The crowns rest in a dental groove that varies in width and depth depending on the TBE. The crown closest to the attached tooth is always the largest and is found in a small resorption pit. As resorption proceeds, the pit expands in volume (cementum and alveolar bone), and the crown increases in size and settles into the pit. Once mature crown size is achieved, the dentine root and cementum portion of the root develop rapidly, the attached tooth is lost and the replacement tooth erupts out of the alveolus. Mosasaurid teeth develop along a 'zig-zag'-shaped movement path: horizontally along the dental groove, down into the alveolus, and up and out of the alveolus prior to attachment to the alveolar wall. At no point in mosasaurid tooth development are the crowns observed in a horizontal position. The mosasaurid dental lamina appears to have been a continuous strip of dental epithelium as it is in other squamates. Mosasaurid tooth attachment is thecodont (histologically and geometrically) not subpleurodont. Most aspects of mosasaurid tooth attachment and ontogeny are autapomorphic for the group.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 687–700.