Balancing function of the masticatory muscles during incisal biting in two murid rodents,Apodemus speciosus and Clethrionomys rufocanus

The functional significance of masticatory muscle direction was estimated using a mechanical model in two murid rodents: the Japanese field mouse (Apodemus speciosus) and the gray red-backed vole (Clethrionomys rufocanus). Theoretical analyses of the data suggest that a balancing mechanism among the muscle forces occurs during incisal power stroke. The activation of the large deep masseter in both murids results in marked tensile separation of two hemimandibles at the flexible mandibular symphysis. Activation of the internal pterygoid decreases this large tensile force at the symphysis more efficiently than other muscles. The lines of action of the deep masseter and internal pterygoid are aligned to produce such a balancing function in both species studied here. The resultant force generated by the deep masseter on both sides is opposite in direction to the reaction force at the lower incisor tip. Therefore, the large deep masseter forms an effective mandibular support mechanism when the reaction forces during biting push the mandible downward. Because of the area of insertion and the line of action, the posterior temporalis appears to have an important role in stabilizing the position of the mandibular condyle in the glenoid fossa during incisal biting. J. Morphol. 236:49–56, 1998. © 1998 Wiley-Liss, Inc.     

Structure and direction of jaw adductor muscles as herbivorous adaptations in <Emphasis Type="Italic">Neotoma mexicana</Emphasis> (Muridae,Rodentia)

The herbivorous adaptations of the jaw adductor muscles in Neotoma mexicana were clarified by a comparative study with an unspecialized relative, Peromyscus maniculatus. In P. maniculatus, the anterior part of the deep masseter arises entirely from the lateral side of an aponeurosis, i.e., superior zygomatic plate aponeurosis, whereas N. mexicana has an additional aponeurosis for this part of the muscle, and the fibers attach on both sides of the superior zygomatic plate aponeurosis. Although the structure of the temporalis muscle is nearly identical in the two genera, a clear aponeurosis of origin occurs only in N. mexicana. These characteristics allow fibrous tissues to be processed with a large occlusal force. The deep masseter, internal pterygoid, and external pterygoid muscles of N. mexicana incline more anterodorsally than those of P. maniculatus. The transverse force component of these muscles relative to whole muscle force is smaller in N. mexicana than in P. maniculatus, with the exception of the internal pterygoid. The anterior part of the temporalis muscle of N. mexicana is specialized to produce occlusal pressure. These findings suggest that in N. mexicana a large anterior force is required to move the heavy mandible, due to the hypsodont molars, against frictional force from food, and that the posterior pull of the temporalis, which adjusts the forward force by the other jaw adductor muscles to a suitable level, need not be large for the mandibular movement.     

New methods of dissecting the masticator space of the rhesus macaque (<Emphasis Type="Italic">Macaca mulatta</Emphasis>)

The aim of our project was to develop a method to examine the deep masticator space of the rhesus macaque (Macaca mulatta) in order to investigate the following particulars of the mandibular nerve: (1) its manner of entry into the masticator space, (2) its branching within spaces/canals in the spheno-temporal bony complex, and (3) the location of its principal divisions with reference to the lateral pterygoid plate and muscle heads. In order to access these structures it was necessary to develop novel lateral and medial approaches. These and the instruments used are described. The proximal branching of the mandibular nerve is described and contrasted with that of the human. The implications of nerve branching in humans with reference to (1) evolution, (2) electromyography and (3) anesthetic control are discussed.     

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.     

Free body analysis,beam mechanics,and finite element modeling of the mandible of Alligator mississippiensis

The mechanical behavior of mammalian mandibles is well‐studied, but a comprehensive biomechanical analysis (incorporating detailed muscle architecture, accurate material properties, and three‐dimensional mechanical behavior) of an extant archosaur mandible has never been carried out. This makes it unclear how closely models of extant and extinct archosaur mandibles reflect reality and prevents comparisons of structure–function relationships in mammalian and archosaur mandibles. We tested hypotheses regarding the mechanical behavior of the mandible of Alligator mississippiensis by analyzing reaction forces and bending, shear, and torsional stress regimes in six models of varying complexity. Models included free body analysis using basic lever arm mechanics, 2D and 3D beam models, and three high‐resolution finite element models of the Alligator mandible, incorporating, respectively, isotropic bone without sutures, anisotropic bone with sutures, and anisotropic bone with sutures and contact between the mandible and the pterygoid flange. Compared with the beam models, the Alligator finite element models exhibited less spatial variability in dorsoventral bending and sagittal shear stress, as well as lower peak values for these stresses, suggesting that Alligator mandibular morphology is in part designed to reduce these stresses during biting. However, the Alligator models exhibited greater variability in the distribution of mediolateral and torsional stresses than the beam models. Incorporating anisotropic bone material properties and sutures into the model reduced dorsoventral and torsional stresses within the mandible, but led to elevated mediolateral stresses. These mediolateral stresses were mitigated by the addition of a pterygoid‐mandibular contact, suggesting important contributions from, and trade‐offs between, material properties and external constraints in Alligator mandible design. Our results suggest that beam modeling does not accurately represent the mechanical behavior of the Alligator mandible, including important performance metrics such as magnitude and orientation of reaction forces, and mediolateral bending and torsional stress distributions. J.Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Reconstruction of the cranial musculature and masticatory function of the Pleistocene panamerican ground sloth Eremotherium laurillardi (Mammalia,Xenarthra, Megatheriidae)

Cranial musculature, dental function and mandibular movement patterns in Eremotherium laurillardi were reconstructed from the examination of crania and dentitions. Size, shape and pattern of muscle divisions were reconstructed from the examination of bony rugosities indicating muscle attachments. Details of masticatory muscle structure and function were based on dissections of the tree sloths Bradypus and Choloepus. Among sloths, masticatory muscles in E. laurillardi demonstrate a different synergist–antagonist pattern, reflecting greater emphasis on mediolateral mandibular movements. Eight cranial character complexes (anterior facial, zygomatic arch, superficial masseter, deep masseter–zygomaticomandibularis, pterygoid, temporal, occipital and occlusal) determined by interrelated contributions of each component made to group functions were identified. An elongate anterior face and predental spout in E. laurillardi allowed protrusion of a long narrow tongue at small degrees of gape, reflecting a probably ancestral xenarthran condition. Gape minimisation, in conjunction with the mediolaterally directed masticatory stroke in E. laurillardi, was a unique solution to increase masticatory efficiency by permitting molariform tooth shearing surfaces to remain in or near occlusion for a greater percentage of each chewing cycle.     

Macaque Pterygoid Muscles: Internal Architecture, Fiber length, and Cross-Sectional Area

Models of mastication require knowledge of fiber lengths and physiological cross-sectional area (PCS), a proxy for muscle force. I dissected 36 medial pterygoid and 36 lateral pterygoid muscles from 30 adult females of 3 macaque species (Macaca fascicularis, M. mulatta, M. nemestrina) using gross and chemical techniques and calculated PCS. These macaques have mechanically similar dietary niches and exhibit no significant difference in muscle architecture or fiber length. Fiber length does not scale with body size (mass) for either total pterygoid muscle or for medial pterygoid muscle mass. However, fiber length scales weakly with lateral pterygoid muscle mass. In each case, differences in PCS among species result from differences in muscle mass not fiber length. Medial pterygoid PCS scales isometrically with body size; larger animals have greater force production capabilities. Medial and lateral pterygoid PCS scale positively allometrically with facial size; individuals with more prognathic faces and taller mandibular corpora have greater PCS, and hence force, values. This positive allometry counters the less efficient positioning of masticatory muscles in longer-faced macaques. PCS is only weakly correlated with bone proxies previously used to estimate muscle force. Thus, predictions of muscle force from bone parameters will entail large margins of error and should be used with caution.     

Diversity and Evolution of the Marsupial Mandibular Angular Process

A medial inflection of the mandibular angular process is present in most marsupials. The few living marsupials that lack this trait either are very specialized forms (e.g., Tarsipes) or show a medial inflection at some point in development that is lost in later ontogenetic stages (cf. Dactylopsila and Phascolarctos). A medially inflected angular process is not present in any known extant or extinct placental (including all Cretaceous taxa that preserve the back of the dentary bone). Some extant placentals with enlarged auditory bullae evolved a medial flange of the angular process as a strategy to increase gape, but this is not homologous to the marsupial condition. We conclude that the medially inflected angular process is a shared derived trait of extant and extinct marsupials. The significant diversity in the form of the medially inflected mandibular angular process in marsupials, documented here for 53 taxa, shows a general relation to dietary adaptations. Herbivores (with well-developed masseter and medial pterygoid muscles) tend to have a shelf-like angular process, while small, insectivorous marsupials generally have a rod-like angular process. A close connection between the angular process and the ectotympanic is maintained during early postnatal development in all marsupials examined, a relation not seen in the placentals examined. A previous hypothesis suggested that the angular process plays a role in hearing in pouch-young Monodelphis. Data on the maturation of the auditory system does not support this hypothesis. Currently there are no data on differences in muscular anatomy or mastication between marsupials and placentals that could serve as a causal explanation for the difference in adult form of the angular process between the two groups.     

Loading patterns and jaw movements during mastication in Macaca fascicularis: a bone-strain, electromyographic, and cineradiographic analysis

Rosette strain gage, electromyography (EMG), and cineradiographic techniques were used to analyze loading patterns and jaw movements during mastication in Macaca fascicularis. The cineradiographic data indicate that macaques generally swallow frequently throughout a chewing sequence, and these swallows are intercalated into a chewing cycle towards the end of a power stroke. The bone strain and jaw movement data indicate that during vigorous mastication the transition between fast close and the power stroke is correlated with a sharp increase in masticatory force, and they also show that in most instances the jaws of macaques are maximally loaded prior to maximum intercuspation, i.e. during phase I (buccal phase) occlusal movements. Moreover, these data indicate that loads during phase II (lingual phase) occlusal movements are ordinarily relatively small. The bone strain data also suggest that the duration of unloading of the jaw during the power stroke of mastication is largely a function of the relaxation time of the jaw adductors. This interpretation is based on the finding that the duration from 100% peak strain to 50% peak strain during unloading closely approximates the half-relaxation time of whole adductor jaw muscles of macaques. The EMG data of the masseter and medial pterygoid muscles have important implications for understanding both the biomechanics of the power stroke and the external forces responsible for the "wishboning" effect that takes place along the mandibular symphysis and corpus during the power stroke of mastication. Although both medial pterygoid muscles reach maximum EMG activity during the power stroke, the activity of the working-side medial pterygoid peaks after the balancing-side medial pterygoid. Associated with the simultaneous increase of force of the working-side medial pterygoid and the decrease of force of the balancing-side medial pterygoid is the persistently high level of EMG activity of the balancing-side deep masseter (posterior portion). This pattern is of considerable significance because the direction of force of both the working-side medial pterygoid and the balancing-side deep masseter are well aligned to aid in driving the working-side lower molars across the upper molars in the medial direction during unilateral mastication.(ABSTRACT TRUNCATED AT 400 WORDS)     

A new specimen of Biseridens qilianicus indicates its phylogenetic position as the most basal anomodont

A new well-preserved basal therapsid skull from the Xidagou Formation, Middle Permian of China, is identified as Biseridens qilianicus. The following synapomorphies distinguish Biseridens as an anomodont and not an eotitanosuchian as previously described: short snout; dorsally elevated zygomatic arch and septomaxilla lacking elongated posterodorsal process between nasal and maxilla. The presence of a differentiated tooth row; denticles on vomer, palatine and pterygoid; contact between tabular and opisthotic; lateral process of transverse flange of pterygoid free of posterior ramus and absence of mandibular foramen exclude it from other anomodonts. Our cladistic analysis indicates Biseridens to be the most basal anomodont, highlights separate Laurasian and Gondwanan basal anomodont clades and suggests that dicynodonts had their origins in the Gondwanan clade. The co-occurrence of the most basal anomodont (Biseridens) together with the most basal therapsid (Raranimus), basal anteosaurid dinocephalians, bolosaurids and dissorophids suggests that the earliest therapsid faunas are from China.     

Description of a new long‐snouted beaked whale from the Late Miocene of Denmark: evolution of suction feeding and sexual dimorphism in the Ziphiidae (Cetacea: Odontoceti)

A new genus and species of Ziphiidae, Dagonodum mojnum gen. nov., sp. nov., from the upper Miocene Gram Formation (c. 9.9–7.2 Ma) represents the first occurrence of the family in Denmark. This long‐snouted ziphiid is characterized by two pairs of mandibular tusks, the Eustachian outlet that approximately levels with the dorsalmost margin of the posterior portion of the involucrum, and the left trapezoid nasal with a posteromedial projection into the frontal. A phylogenetic analysis including 25 species and 69 characters was conducted. Dagonodum mojnum is placed in a basal ziphiid clade as the sister taxon of Messapicetus. The specimen is probably a male, because it has enlarged tusks. Alternatively, females could also be involved in fights and develop erupted tusks as in the extant Berardius. Although less well supported, this interpretation proposes that aggressive interactions were not restricted to males in stem‐ziphiids. With a thickened thyrohyal and the presence of a precoronoid crest, D. mojnum was able to use suction feeding, but was less specialized to it compared to extant ziphiids. The elongated neck of D. mojnum less optimized to perform deep dives, and the shallow depth at which the Gram Formation was deposited corroborates the hypothesis that at least part of the stem‐ziphiids were not regular deep divers.     

The Xujiayao 14 Mandibular Ramus and Pleistocene Homo Mandibular Variation

The earlier Late Pleistocene mandibular ramus from Xujiayao (northern China) preserves traits that vary distributionally among western Old World Pleistocene Homo samples and between Early/Middle Pleistocene archaic humans and Late Pleistocene modern humans in eastern Eurasia. Xujiayao 14 presents a lateral mandibular notch crest, an open mandibular foramen, a wide ramus, an asymmetrical mandibular notch, an enlarged superior medial pterygoid tubercle, (probably) a retromolar space, and gonial eversion, as well as an unusual depression in the planum triangulare. The first two traits appear ancestral for Later Pleistocene and recent Homo and are dominant among modern humans. The second two traits largely separate Xujiayao 14 and archaic Homo from modern humans. The next two traits are found in the highest frequency among the Neandertals, although gonial eversion contrasts with Late Pleistocene Neandertals. Xujiayao 14, in the context of Pleistocene and recent Homo samples and the other Xujiayao human remains, therefore provides a morphological mosaic, highlighting regional variation through the Pleistocene.     

Biomechanics of cranial kinesis in birds: Testing linkage models in the white-throated sparrow (Zonotrichia albicollis)

Cranial kinesis in sparrows refers to the rotation of the upper jaw around its kinetic joint with the braincase. Avian jaw mechanics may involve the coupled motions of upper and lower jaws, in which the postorbital ligament transfers forces from the lower jaw, through the quadrate, pterygoid, and jugal bones, to the upper jaw. Alternatively, jaw motions may be uncoupled, with the upper jaw moving independently of the lower jaw. We tested hypotheses of cranial kinesis through the use of quantitative computer models. We present a biomechanical model of avian jaw kinetics that predicts the motions of the jaws under assumptions of both a coupled and an uncoupled mechanism. In addition, the model predicts jaw motions under conditions of force transfer by either the jugal or the pterygoid bones. Thus four alternative models may be tested using the proposed model (coupled jugal, coupled pterygoid, uncoupled jugal, uncoupled pterygoid). All models are based on the mechanics of four-bar linkages and lever systems and use morphometric data on cranial structure as the basis for predicting cranial movements. Predictions of cranial motions are tested by comparison to kinematics of white-throated sparrows (Zonotrichia albicollis) during singing. The predicted relations between jaw motions for the coupled model are significantly different from video observations. We conclude that the upper and lower jaws are not coupled in white-throated sparrows. The range of jaw motions during song is consistent with a model in which independent contractions of upper and lower jaw muscles control beak motion. © 1996 Wiley-Liss, Inc.     

Mandibular movement and muscle activity during mastication in the guinea pig (Cavia porcellus)

Optoelectronic analysis of mandibular movement and electromyography (EMG) of masticatory muscles in Cavia porcellus indicate bilateral, unilateral, and gnawing cycles. During bilateral and unilateral cycles, the mandibular tip moves forward, lateral, and down during the lingual phase of the power stroke to bring the teeth into occlusion. EMG activity is generally asymmetric, with the exception of activity of the temporalis muscle during bilateral cycles. During gnawing cycles, the mandible moves in an anteroposterior direction that is opposite that during bilateral and unilateral chew cycles. Bilateral and unilateral cycles of pellets were significantly longer than carrot. With the exception of the width of bilateral cycles, the magnitude of cycle width, length, and height during the mastication of carrots was greater than that during the mastication of pellets. Significant differences exist between EMG durations during mastication of pellets and carrots. The lateral pterygoid displays continuous activity during gnawing cycles. Significant differences also exist in the durations of EMG activity between the working and balancing side during all three cycle types. High level activity of balancing side temporalis and anterior belly of digastric (ABD) during bilateral cycles occurs during rotation and depression of the mandible during the power stroke. The temporalis apparently provides a ?braking”? or compensatory role during closing and power strokes. Differences between Cavia masticatory patterns and those shown by Rattus and Mesocricetus are apparently due to differences in dental morphology, occlusal relationships, and, possibly, the poorly developed temporalis in Cavia. The large number and wide diversity of rodent groups afford students of mammalian mastication an opportunity to investigate and compare different masticatory specializations.     

Quantitative assay of electromyograms during mastication in domestic cats (Felis catus)

Mastication has been studied by cinematography with synchronized electromyography (computer quantified and analyzed), while unanesthetized, freely feeding cats (Felis catus) were reducing equivalent-sized chunks of raw and cooked beef and cooked chicken. Cats reduce food on one side at a time, and their chewing cycles show both horizontal and anteroposterior deflections. Food objects are shifted from side to side by lateral jerks of the head and movements of the tongue. During the opening phase, the lower jaw is rotated relatively straight downward, and the digastric muscles are active in bilateral symmetry. Near the end of opening, the head jerks upward, both zygomaticomandibulares start to fire, and opening acceleration of the mandible decreases. Closing starts with horizontal displacement of the mandibular canines toward the working side, accompanied by asymmetrical activities from the working side deep temporalis and the balancing side medial pterygoid, as well as a downward jerk of the head. As closing proceeds, the mandibular canines remain near the working side and the working side zygomaticomandibularis and deep masseter are very active. Near the end of closing, the mandibular canine on the working side moves toward the midline, and adductors, digastrics, and lateral pterygoids of both sides are active. The adductors of the working side are generally more active than those of the balancing side. During a reduction sequence, the number and shape of the masticatory cycles, as well as movements of the head, during a reduction sequence are affected significantly by food type. As reduction proceeds, the duration of bite and the muscular activity (as characterized by number and amplitude of spikes) change significantly among muscles of the working and balancing sides. The adductors of the working side are generally most active when cats chew raw beef, less for cooked beef, and least for cooked chicken. In general, the adductor activity reflects food consistency, whereas that of the digastrics and lateral pterygoids reflects more the vertical and lateral displacements of the mandible. Statistical analysis documents that the methods of electrode insertion and test give repeatable results for particular sites in different animals. Thus, it should be possible to compare these results with those produced while other mammalas are masticating.     

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.     

Biomechanical scaling of mandibular dimensions in New World Monkeys

Previous studies show that folivorous Old World monkeys have shorter, deeper mandibles and shorter, wider condyles than frugivorous ones. These morphologies have been related to leaf mastication in colobines and ingestion of large, tough fruits in cercopithecines. This study examines New World monkeys in order to determine whether they exhibit similar adaptations to diet. New World monkeys have relatively long, transversely thin mandibles and somewhat deep mandibles and narrow condyles. Except for their deep mandibles, folivorous New World monkeys (i.e., Alouatta) do not exhibit the mandibular and condylar specializations typical of cercopithecid folivores. Reliance on comparatively nonfibrous foods plus alterations in masticatory muscle ratios among New World monkeys partially accounts for observed differences between folivorous New and Old World monkeys. In addition, adaptations for howling in Alouatta appear to have a significant effect on mandibular morphology. A biomechanical interpretation of craniofacial scaling patterns suggests that the mandibles of New World monkeys are subjected to lower condylar loads and considerably less twisting of the mandibular corpus than those of comparable Old World monkeys.     

A comparative study of incisor procumbency and mandibular morphology in vampire bats

The three species of vampire bats (Phyllostomidae: Desmodontinae), Desmodus rotundus, Diaemus youngi, and Diphylla ecaudata, are the only mammals that obtain all nutrition from vertebrate blood (sanguinivory). Because of the unique challenges of this dietary niche, vampire bats possess a suite of behavioral, physiological, and morphological specializations. Morphological specializations include a dentition characterized by small, bladelike, non‐occlusive cheek teeth, large canines, and extremely large, procumbent, sickle‐shaped upper central incisors. The tips of these incisors rest in cuplike pits in the mandible behind the lower incisors (mandibular pits). Here, we use microCT scanning and high‐resolution radiography to describe the morphology of the mandible and anterior dentition in vampire bats, focusing on the relationship between symphyseal fusion, mandibular pit size, incisor size, and procumbency. In Desmodus and Diaemus, highly procumbent upper incisors are associated with relatively small mandibular pits, an unfused mandibular symphysis with substantial bony interdigitations linking the dentaries, and a diastema between the lower central incisors that helps to facilitate the lapping of blood from a wound. In Diphylla, less procumbent upper incisors are associated with relatively large mandibular pits, a completely fused mandibular symphysis, and a continuous lower toothrow lacking a central diastema. We hypothesize that symphyseal morphology and the presence or absence of the diastema are associated with the angle of upper incisor procumbency and mandibular pit development, and that spatial constraints influence the morphology of the symphysis. Finally, this morphological variation suggests that Diphylla utilizes a different feeding strategy as compared to Desmodus and Diaemus, possibly resulting from the functional demands of specialization on avian, rather than mammalian, blood. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Osteology and cranial musculature of Caiman latirostris (crocodylia: Alligatoridae)

Caiman latirostris Daudin is one of the extant species of Caimaninae alligatorids characterized taxonomically only by external morphological features. In the present contribution, we describe the cranial osteology and myology of this species and its morphological variation. Several skull dissections and comparisons with other caimans were made. Although jaw muscles of living crocodiles show the same general “Bauplan” and alligatorids seem to have a similar cranial musculature pattern, we describe some morphological variations (e.g., in C. latirostris the superficial portion of the M. adductor mandibulae externus did not reach the postorbital; the M. adductor mandibulae internus pars pterygoideus dorsalis did not reach the pterygoid and lacrimal and contrary to the case of C. crocodilus the M. adductor mandibulae internus pars pterygoideus ventralis attaches to the posterodorsal surface of the pterygoid and the pterygoid aponeurosis, without contacting the dorsal and ventral surface of the pterygoid margin; the M. intermandibularis is attached to the anterior half of the splenial and posteriorly inserts medially by a medial raphe that serves as attachment zone for M. constrictor colli, and the M. constrictor colli profundus presents a medial notch in its anterior margin). In addition, the skull of C. latirostris differs from that of other caimans and possesses several characters that are potential diagnostic features of this species (e.g., outline of glenoid cavity in dorsal view, extension of the rostral ridges, and occlusion of the first dentary tooth). Nevertheless, these characters should be analyzed within the phylogenetic context of the Caimaninae to evaluate its evolutionary implications for the history of the group. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Development and diversity of the suspensorium of trichomycterids and comparison with loricarioids (Teleostei: Siluriformes)

Studies of ontogenetic series of trichomycterids and other catfishes reveal that the suspensorium of siluroids is highly specialized; several synapomorphies separate siluroids from other teleosts. In siluroids, the palatoquadrate is divided into pars autopalatina and pars pterygoquadrata and both are usually connected by the autopaiatine-metapterygoid ligament. The pterygoquadrate is broadly joined to the dorsal limb of the hyoid arch, forming a cartilaginous hyomandibular-symplectic-pterygoquadrate plate in early ontogeny. This produces a special alignment of the hyomandibula and quadrate which is characteristic of siluroids. A symplectic bone is absent. The interhyal is absent in trichomycterids and astroblepids. Dorsal and ventral limbs of the hyoid arch are connected by a ligament. A rudimentary interhyal and this ligament are present in primitive siluroids such as diplomystids and nematogenyids as well as loricariids. The metapterygoid arises as an anterior ossification of the pars pterygoquadrata in siluroids. The formation and position of the metapterygoid exhibit two patterns: (1) the metapterygoid develops as an ossification of a cartilaginous projection positioned between the future hyomandibula and quadrate in primitive catfishes (e.g., Diplomystes) as well as in Nematogenys, callichthyids, loricariids, and astroblepids; (2) the metapterygoid arises as an ossification of the cartilaginous projection (pterygoid process) positioned just above the articular facet of the quadrate for the lower jaw. An ossified anterior chondral pterygoid process of the complex quadrate is present in trichomycterids, whereas the process is absent (simple quadrate) in catfishes such as diplomystids, nematogenyids, callichthyids, and loricariids. The anterior membranous process of the quadrate of Astroblepus is non-homologous with the chondral pterygoid process of trichomycterids; both structures arose independently within the loricarioids. Despite topological relationships, the origin and development of bones reveal the presence of a chondral hyomandibula which develops a large meinbranous outgrowth during ontogeny and a chondral metapterygoid in trichomycterids. The presence of a compound hyomandibula + metapterygoid or a compound metapterygoid + ectopterygoid + entopterygoid have no developmental support in trichomycterines or other siluroids. The “entopterygoid” of Nematogenys and Diplomystes arises as an ossification of a ligament. The dermal entopterygoid of other ostariophysans and the “entopterygoid” are homologous. An ectopterygoid or tendon bone “ectopterygoid” is absent in loricarioids. The suspensorium is an important structural system which has significant evolutionary transformations which characterize loricarioid subgroups; however, no character, of the suspensorium supports the monophyly of the loricarioids.