Anatomy and Ultrastructure of Ventral Pharyngeal Organs and their Phylogenetic Importance in Polychaeta (Annelida). IV. The Pharynx and Jaws of the Dorvilleidae

An anatomical and ultrastructural investigation of the ventral pharyngeal organ, jaws and replacement of jaws was carried out in Ophryotrocha gracilis and Protodorvillea kefersteini (Dorvilleidae). The pharynx exhibits the following features: jaw apparatus present, consisting of paired mandibles and rows of maxillary plates, the latter are fused to form a single piece; cuticular jaws electron-dense, in P. kefersteini with collagen fibres; muscle bulbus solid, composed of muscle cells only; parallel running myofilaments, centrally located mitochondria and nuclei, bulbus epithelium containing the mandibles and gland cells, maxillary plates lying on folds corresponding to a tongue-like organ, connected with mandibles by longitudinal investing muscles; numerous gland cells not united to distinct salivary glands. Development of jaw replacements occurs in epithelial cavities beside the functional maxillae. Shape of maxillary plates is preformed by microvilli carrying cell processes. Maxilloblasts change their shape during the development. Synapomorphic structures occurring in ventral pharyngeal organs of other species outside the Eunicea are not present and even the closely related Dinophilidae exhibit a completely different pharyngeal organ. Therefore, convergent evolution of these organs is the most probable explanation. These findings do not agree with the hypothesis of the homology of the ventral pharyngeal organs in the Polychaeta.     

Trigonognathus kabeyai,a new genus and species of the squalid sharks from Japan

Two specimens of the peculiar squalid shark,Trigonognathus kabeyai gen. et sp. nov., were collected from the coastal waters of Wakayama and Tokushima, Japan, by bottom trawl at depths of 330 and 360 meters. Shape of teeth similar in both jaws; slender, unicuspid, canine-like, without any cusplets or serrations, with weak thin fold on both lingual and labial sides in anterior teeth on both jaws; tooth at symphysis of each jaw longest. Interspace between teeth very wide. Both jaws triangular in shape. Most of dermal denticles on body and head roughly rhombic, swollen very much near central part, with about 10–40 facets on the dorsal surface of its crown. Preoral snout length very short. Many small organs considered to be photophores present mainly on ventral surfaces of head and body.     

Vertebrate dentitions at the origin of jaws: when and how pattern evolved

New evidence shows that teeth evolved with a greater degree of independence from jaws than previously considered. Pharyngeal denticles occur in jawless fish and also in early gnathostomes and precede jaw teeth in phylogeny. Many of these denticles form joined polarized sets on each branchial arch; these resemble whorl-shaped tooth sets on the jaws of stem and crown gnathostomes and are proposed as homologous units. Therefore, the source of patterning of these pharyngeal denticle and tooth sets is conserved from jawless conditions. It is proposed that developmental regulatory systems, responsible for all such tooth patterns on the jaws, are co-opted from the pharyngeal region and not from the skin as classically understood. This strongly implicates embryonic endoderm as opposed to ectoderm in the genetic control of dentition patterning. New interpretations of ontogenetic data on patterning dentitions of extant sharks are proposed, together with those of osteichthyan fish. Two entirely fossil groups, placoderms and acanthodians, at the base of gnathostome phylogeny are reassessed on the basis of a new model. It is concluded that within stem group and crown group gnathostomes several different strategies, unique to each taxon, were adopted to produce different developmental models of dentition patterning from pharyngeal denticles. One shared developmental pattern is that of initiation from primordial tooth sites, independently in each dentate zone of the jaws. The new model is proposed as a framework for data on evolutionary developmental genetics.     

A revised hypothesis on the evolutionary origin of the vertebrate dentition

Despite claims to the contrary, the evolutionary origin of teeth has not been definitely established. The classical ‘outside in’ theory stating that teeth derive from odontodes that invaded the oral cavity in conjunction with the origin of jaws has been challenged by an alternative, ‘inside out’, hypothesis suggesting that teeth evolved from pharyngeal denticles, as endodermal derivatives, prior to the origin of jaws. We propose a third scenario, a revised ‘outside in’ hypothesis ( Huysseune et al., 2009 ). Our hypothesis is consistent with the current data and avoids speculations about convergent tooth evolution. We suggest that teeth may indeed have arisen before the origin of jaws, a pillar of the ‘inside out’ hypothesis, but not from the endodermally lined posterior pharynx. Rather, teeth would have been the result of competent, odontode‐forming ectoderm invading the oropharyngeal cavity through the mouth as well as through the gill slits, interacting with neural‐crest derived mesenchyme. Arguments in support of this hypothesis are: (i) the observation that pharyngeal teeth are present only in species known to possess gill slits, and disappear from the pharyngeal region in early tetrapods concomitant with the closure of gill slits; (ii) the assumption that endoderm alone, together with neural crest, cannot form teeth; (iii) observations on pharyngeal tooth and gill slit formation in extant species; (iv) the observation that the dental lamina (sensu Reif, 1982 ) is not a prerequisite for tooth formation; (v) evidence that patterning does not distinguish pharyngeal from skin denticles, and (vi) the observation on zebrafish mutants affected in the dermal skeleton. This ‘modified outside in’ hypothesis can be tested both on paleontological data (it predicts a correlation of the presence of pharyngeal teeth and of gill slits), and on developmental data in extant species (it predicts the necessity of an ectodermal signal to make [pharyngeal] teeth).     

Jaw growth and replacement in<Emphasis Type="Italic"> Ophryotrocha labronica</Emphasis> (Polychaeta,Dorvilleidae)

Ophryotrocha labronica, as typical for Eunicida, has a complex jaw apparatus consisting of ventral mandibles and dorsal maxillae. Mandibles are not replaced but are retained throughout life. Larval mandibles have adult-sized cutting plates but their proximal shafts lengthen and enlarge as the worm grows. The maxillary apparatus of O. labronica undergoes three moults or replacements. The initial, or larval maxillae, consisting of two paired basal plates and two paired free denticles, develop in the unreleased larvae. They are replaced in the 5-setiger juvenile by the P1-maxillae consisting of falcate forceps and six denticles. The second moult occurs in the 8- to 9-setiger juveniles and results in the P2-maxillae with bidentate forceps and seven denticles, and the third and final moult results in the K-maxillae and seven denticles. The K-maxillae develop in 9- to 12-setiger males and 13- to 15-setiger females and are not replaced but enlarge proximally. Thus the K-forceps can be traced back through the P2-forceps, P1-forceps, to the larval basal plates, indicating the apomorphic state of the K-forceps. Three pulp cavities, separated by darker fusion lines are visible in weakly sclerotised young K-forceps suggesting the fusion of three separate elements. It is concluded that the Ophryotrocha forceps are homologous to the superior and probably inferior basal plates of other dorvilleids. The internal structure of the Ophryotrocha forceps demonstrates that they are not homologous to the labidognath maxilla I as has been suggested.     

The structure and function of a muscle articulation‐type jaw joint of a polychaete worm

The arrangement of the musculature and the fibers of the extracellular matrix (ECM) in the flexible jaw joint of the sandworm Alitta virens (Annelida, Polychaeta) was studied using dissection and histology. The jaws are capable of a wide range of motions principally related to defense and feeding. The left and right jaws are embedded in and moved by a compact pharyngeal bulb of muscle and ECM that also forms the mouth and esophagus. Eight pharyngeal bulbs were removed and dissected to document gross anatomical features or preserved and embedded in plastic for sectioning in multiple planes. The sections were stained with toluidine blue and basic fuchsin to differentiate muscle and ECM. The sections were then digitized and used to develop a three‐dimensional computer illustration. We hypothesize that the muscle and fibers in the ECM are arranged as a muscular hydrostat to support the movement of the jaws. Four specimens were recorded using a digital video camera and a tank with an angled mirror to record lateral and ventral views of jaw movements during locomotion and biting associated with burrow guarding and feeding. Frame by frame kinematic analysis of this video showed that the jaws move symmetrically in a roughly horizontal plane. Although the angle between the jaws increases and then decreases after maximum gape has been reached, the jaws also translate relative to each other such that the axis of rotation is not fixed. Together, these functional morphological and behavioral data identify the jaw mechanism as a flexible joint known as a muscle articulation. As muscle articulations have been previously described only in the beaks of cephalopods and flatworms, this study implies that this type of joint is more common and important than previously recognized. J. Morphol. 276:403–414, 2015. © 2014 Wiley Periodicals, Inc.     

南鳅属鱼类一新种及南鳅属评述

记述了南鳅属鱼类一新种－－异颌南鳅Ｓｃｈｉｓｔｕｒａ ｈｅｔｅｒｏｇｎａｔｈｏｓ ｓｐ．ｎｏｖ．,用作新种描述的模式标本共７属,其中正模标本１属,体长８０．０ｍｍ,编号８８１０２４０;副模标本６属,体长９５．０－１０５．０ｍｍ,编号８８１０２４１、８８１０２４１、８８１０１５０￣８８１０１５３。１９８８年采自云南省勐腊县的那着和曼庄,属澜沧江下游支流南腊河。所有模式标本均保存于中国科学院水生生物研     

Tooth growth and replacement in Ctenolucius hujeta, a neotropical characoid fish

The predaceous neotropical characoid fish Ctenolucius has an essentially homodont dentition, the number of teeth increasing linearly with age. The basic manner of tooth replacement suggests that Ctenolucius is a primitive characoid. Tooth replacement continues throughout life and is similar to that of tetrapods, involving replacement waves which pass from the back to the front of the jaws. The waves containing the greatest number of teeth are found just anterior to the middle of the jaws. In the upper jaw the increase in the number of teeth is restricted to the anterior portion (premaxillary) whereas the number on the posterior part (maxillary) remains constant. In specimens measuring from 68–230 mm in standard length the posterior portion of the upper jaw doubles in length whereas the anterior portion triples. It is suggested that the area immediately anterior to the middle of the jaw, where replacement waves are longest, is where most of the increase in tooth numbers occurs. During growth of the teeth the absolute height is always greater than the absolute width as the shape changes. The final shape of the recurved conical teeth is determined only in the last stages of tooth formation when the main axis of growth abruptly changes.     

Mechanics of biting in great white and sandtiger sharks

Although a strong correlation between jaw mechanics and prey selection has been demonstrated in bony fishes (Osteichthyes), how jaw mechanics influence feeding performance in cartilaginous fishes (Chondrichthyes) remains unknown. Hence, tooth shape has been regarded as a primary predictor of feeding behavior in sharks. Here we apply Finite Element Analysis (FEA) to examine form and function in the jaws of two threatened shark species, the great white (Carcharodon carcharias) and the sandtiger (Carcharias taurus). These species possess characteristic tooth shapes believed to reflect dietary preferences. We show that the jaws of sandtigers and great whites are adapted for rapid closure and generation of maximum bite force, respectively, and that these functional differences are consistent with diet and dentition. Our results suggest that in both taxa, insertion of jaw adductor muscles on a central tendon functions to straighten and sustain muscle fibers to nearly orthogonal insertion angles as the mouth opens. We argue that this jaw muscle arrangement allows high bite forces to be maintained across a wider range of gape angles than observed in mammalian models. Finally, our data suggest that the jaws of sub-adult great whites are mechanically vulnerable when handling large prey. In addition to ontogenetic changes in dentition, further mineralization of the jaws may be required to effectively feed on marine mammals. Our study is the first comparative FEA of the jaws for any fish species. Results highlight the potential of FEA for testing previously intractable questions regarding feeding mechanisms in sharks and other vertebrates.     

Jaw mechanism and dental function in the late cretaceous basal eusuchian Iharkutosuchus

Iharkutosuchus makadii is a basal eusuchian crocodylian with multicusped teeth discovered from the Upper Cretaceous of Hungary. Skull and dentition morphology indicates an active food processing for this crocodylian. First among crocodylians, a combination of different analyses, including cranial adductor muscle reconstruction, tooth wear pattern, and enamel microstructure studies, is applied here to support this hypothesis. Data provide unambiguous evidence for significant dental occlusion that was a result of a unique, transverse mandibular movement. Reconstruction of the jaw adductors demonstrates strong muscles responsible for slow but active jaw closure as the motor of transverse jaw movement; nevertheless muscles producing rapid jaw closure were reduced. Macrowear orientations show a dominantly transverse movement of the mandibles completed by a slight anteroposterior component. Along with quadrate morphology, macrowear further indicates that this motion was accomplished by alternate rotation of the mandibles about the quadrate condyles. Dental morphology and wear patterns suggest two types of power stroke: a slicing–crushing stroke associated dominantly with anterior tooth–food–tooth contact (with a low degree of transverse mandibular movement) during in the early stage of mastication, and a grinding stroke with significant posterior tooth–tooth contact and a dynamic transverse movement occurring later. The patterns of microwear show a diverse diet for Iharkutosuchus including both soft and hard items. This is also supported by the microstructure of the thick, wrinkled enamel built up mostly by poorly developed columnar units. Based on wear patterns, ontogenetic variation in feeding habits of Iharkutosuchus is also recognized. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

Do sharks exhibit heterodonty by tooth position and over ontogeny? A comparison using elliptic Fourier analysis

Tooth morphology is often used to inform the feeding ecology of an organism as these structures are important to procure and process dietary resources. In sharks, differences in morphology may facilitate the capture and handling of prey with different physical properties. However, few studies have investigated differences in tooth morphology over ontogeny, throughout the jaws of a single species, or among species at multiple tooth positions. Bull (Carcharhinus leucas), blacktip (Carcharhinus limbatus), and bonnethead sharks (Sphyrna tiburo) are coastal predators that exhibit ontogenetic dietary shifts, but differ in their feeding ecologies. This study measured tooth morphology at six positions along the upper and lower jaws of each species using elliptic Fourier analysis to make comparisons within and among species over their ontogeny. Significant ontogenetic differences were detected at four of the six tooth positions in bull sharks, but only the posterior position on the lower jaw appeared to exhibit a functionally relevant shift in morphology. No ontogenetic changes in morphology were detected in blacktip or bonnethead sharks. Intraspecific comparisons found that most tooth positions significantly differed from one another across all species, but heterodonty was greatest in bull sharks. Additionally, interspecific comparisons found differences among all species at each tooth position except between bull and blacktip sharks at two positions. These morphological patterns within and among species may have implications for prey handling efficiency, as well as in providing insight for paleoichthyology studies and reevaluating heterodonty in sharks.     

A quantitative theory of expected volume changes of the mouth during feeding in teleost fishes

The mechanism of mouth expansion in fish, consisting of jaws, suspensoria (j) and hyoids (h) has been modelled by a four-bar isosceles linkage. This model provides insight into limitations and demands of the expansion system used in feeding, as it can be optimized with regard to maximum mouth volume increase. The optimum length ratio of hyoids and jaws was found to be h/j = 0–7. This optimum is modified by mouth bottom depression, jaw protrusion and swimming.
To expand the mouth, at least two forces are required; one exerted by the sternohyoid and ventral body muscles, the other by the epaxial muscles through transmission in the quadrato-articular joints. (Data from EMG experiments confirm the synchronous activity of these muscle groups.) Force transmission and mouth volume increase are constraining quantities, which can be compromised. This leads to a model of the initial mouth shape which is actually found in many 'generalized' fishes, and to demands concerning volume and physiological cross-sectional area of the muscles involved.
Options for specific relative lengths of jaws and hyoids (h/j-ratios) are, for various fish species, compared with model predictions. The applicability of the model approach is shown by the obtained results.     

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.     

How to recognize in situ fossil cephalopods: evidence from experiments with modern Nautilus

Field and flume experiments with modern Nautilus pompilius establish two prerequisites to recognize in situ preservation of fossil cephalopod shells (soft parts were within body chamber in situ at the time of fossilization): occurrence of the upper jaw within the body chamber and the position of jaws within the body chamber. Morphology of shells and jaws in modern and fossil nautiloids is so similar that these prerequisites can be applied for fossil nautiloids and provide implications for ammonoids. The upper jaws of Nautilus start to move at a water velocity of > 0.2 m/s, when the shells are reoriented with the aperture downstream; jaws are therefore unlikely to be secondarily deposited near the shell aperture by bottom currents. The lower jaws, moved at the velocity of > 0.1 m/s, can be deposited around the shell aperture by weak current (0.1–0.2 m/s in velocity), but never enter the inside of body chamber. Neither jaw is likely to be separately and selectively displaced from the inside of the body chamber through scavenging of the soft parts by burrowing infaunal animals. An upper jaw preserved inside the body chamber, together with a lower jaw, is thus a reliable indicator of in situ preservation; a sole lower jaw preserved around the shell aperture is likely to be secondarily deposited. Sedimentary structures inferring rapid burial events and jaw size are useful as additional evidence. Smaller jaws were more likely to be displaced from the body chamber by scavenging by infaunal animals after in situ burial, so that smaller jaws preserved within the body chamber suggest less scavenging. These findings are crucial to interpreting the taphonomic history and palaeo-ecology of fossil cephalopods.     

The tadpole of Scinax skuki (Anura: Hylidae) from the type locality,with a description of its larval skeleton

Herein, we provide external and internal morphological data of Scinax skuki tadpoles from its type locality. The benthic tadpole of S. skuki has eyes and nostrils positioned dorsally, vent tube dextral and reaching the free margin of the ventral fin, oral disk ventral with posterior margin concave when partially closed, labial tooth row formula 2/3, and the presence of nonpigmented spurs behind the lower jaw. These characters, together with the absence of a tectum parietale, and the shapes of the pars articularis quadrati and suprarostral, are useful for species identification and may be informative for systematic purposes.     

Fine morphology of the jaw apparatus of Puncturella noachina (Fissurellidae,Vetigastropoda)

Jaws of various kinds occur in virtually all groups of Mollusca, except for Polyplacophora and Bivalvia. Molluscan jaws are formed by the buccal epithelium and either constitute a single plate, a paired formation or a serial structure. Buccal ectodermal structures in gastropods are rather different. They can be nonrenewable or having final growth, like the hooks in Clione (Gastropoda, Gymnosomata). In this case, they are formed by a single cell. Conversely, they can be renewable during the entire life span and in this case they are formed by a set of cells, like the formation of the radula. The fine structure of the jaws was studied in the gastropod Puncturella noachina. The jaw is situated in the buccal cavity and consists of paired elongated cuticular plates. On the anterior edge of each cuticular plate there are numerous longitudinally oriented rodlets disposed over the entire jaw surface and immersed into a cuticular matrix. The jaw can be divided into four zones situated successively toward the anterior edge: 1) the posterior area: the zone of formation of the thick cuticle covering the entire jaw and forming the electron‐dense outer layer of the jaw plate; 2) the zone of rodlet formation; 3) the zone of rodlet arrangement; and 4) the anterior zone: the free scraping edge of the plate, or the erosion zone. In the general pattern of jaw formation, Puncturella noachina resembles Testudinalia tessulata (Patellogastropoda) studied previously. The basis of the jaw is a cuticular plate formed by the activity of the strongly developed microvillar apparatus of the gnathoepithelium. However, the mechanism of renewal of the jaw anterior part in P. noachina is much more complex as its scraping edge consists not just of a thick cuticular matrix rather than of a system of denticles being the projecting endings of rodlets. J. Morphol. 275:775–787, 2014. © 2014 Wiley Periodicals, Inc.     

Morphology and mechanics of the teeth and jaws of white-spotted bamboo sharks (Chiloscyllium plagiosum)

The teeth of white-spotted bamboo sharks (Chiloscyllium plagiosum) are used to clutch soft-bodied prey and crush hard prey; however, the dual function is not evident from tooth morphology alone. Teeth exhibit characteristics that are in agreement with a clutching-type tooth morphology that is well suited for grasping and holding soft-bodied prey, but not for crushing hard prey. The dual role of this single tooth morphology is facilitated by features of the dental ligament and jaw joint. Tooth attachment is flexible and elastic, allowing movement in both sagittal and frontal planes. During prey capture spike-like tooth cusps pierce the flesh of soft prey, thereby preventing escape. When processing prey harder than the teeth can pierce the teeth passively depress, rotating inward towards the oral cavity such that the broader labial faces of the teeth are nearly parallel to the surface of the jaws and form a crushing surface. Movement into the depressed position increases the tooth surface area contacting prey and decreases the total stress applied to the tooth, thereby decreasing the risk of structural failure. This action is aided by a jaw joint that is ventrally offset from the occlusal planes of the jaws. The offset joint position allows many teeth to contact prey simultaneously and orients force vectors at contact points between the jaws and prey in a manner that shears or rolls prey between the jaws during a bite, thus, aiding in processing while reducing forward slip of hard prey from the mouth. Together the teeth, dental ligament, and jaws form an integrated system that may be beneficial to the feeding ecology of C. plagiosum, allowing for a diet that includes prey of varying hardness and elusiveness.     

Morphological adaptation to diet in platyrrhine primates

Morphological features of the jaws and teeth are examined in eight species of platyrrhine monkeys that coexist in the Suriname rainforest. Z-scores calculated from geometric predictions for several features of the feeding apparatus thought to have some functional significance (e. g., tooth dimensions, jaw robusticity, leverage of primary jaw elevators) are compared to a profile of the naturalistic dietary behavior of these species (i. e., proportions of fruit mesocarp, seeds, leaves, and fauna eaten). Several features are found exclusively in those platyrrhines whose dietary preferences are the most limited. Such specializations appear to be associated with a particular protein source exploited by a species to supplement a largely frugivorous diet. Ateles paniscus, which feeds primarily on the mesocarp of ripe fruit, has an adaptive morphology that emphasizes broad incisors. Chiropotes satanas (and to a slightly lesser extent, Pithecia pithecia) is a frugivore/seed predator with large upper and lower canines and a robust mandible. The frugivore/folivore Alouatta seniculus has a relatively large total molar area and effective mandibular condyle height. In all four of these strictly vegetarian species, the leverage of the masseter muscle is greater than that of temporalis. Of the omnivorous species, Cebus apella and C. nigrivittatus exploit both fauna and seeds for protein and exhibit an array of many of the above features, such as large teeth and thick mandibles. Saimiri sciureus, not particularly known for seed predation, departs from Cebus in having less robust canines and a more gracile mandible. All three cebid omnivores have a temporalis with greater leverage than the masseter, indicating a requirement for resisting anteriorly directed forces, for example, using the jaws for vigorous foraging. The lack of any enlarged features, other than incisors, in the omnivorous Saguinus midas may be attributable to the functional constraints of small body size. Because the small size of the gape limits the size of the food parcel ingested, a requirement to enlarge other dentomandibular structures for trituration is alleviated.