The development of the trigeminal jaw adductor musculature in the grass snake Natrix natrix

The ontogenetic development of the jaw adductor musculature in Natrix natrix (L.) is described in detail and related to patterns of ossification in the skull. A comparison with the development of the jaw adductors in the lizard Podarcis sicula reveals some interesting differences of dynamics in the developing head. The problem of the establishment of topological relations of similarity (homology) in developing systems is discussed.
The homologies of the external jaw adductor compartments in lizards and snakes are revised. Early ontogenetic divergence explains the reversal of fibre direction in the anterior portion of the external adductor of snakes, and renders the homologization of that fibre bundle with part of the external adductor in lizards impossible.     

The structure and development of the jaw adductor musculature in the turtle Chelydra serpentina

The investigation of the development of the trigeminal jaw adductor musculature in the turtle Chelydra serpentina documents the early aggregation of muscle rudiments around the innervating nerve branches, probably a consequence of inductive interaction. This may explain the early continuity of the intramandibularis with the intermandibularis muscle. Several aspects of muscle development differ in the turtle as compared to lizards. These differences highlight the fact that conjectures of homology, based on a static topographical correspondence of adult structures, cannot capture the dynamics of the developmental process. The intramandibularis muscle of turtles, comparable to that of crocodiles, represents a plesiomorphous structure which is not homologous to the intramandibularis muscle of lacertoid lizards, a derived feature of the Lacertoidea. A derived feature of the chelonian jaw adductor musculature is the posterodorsal expansion of the external adductor along a supraoccipital crest, developing according to a pattern of Haeckelian recapitulation. Muscle development serves to corroborate the concept of a monophyletic Eureptilia, including diapsids and synapsids, as opposed to the (paraphyletic) Anapsida. The impact of the differentiation of the external adductor into a pulley system on cranial kinesis is analysed in biomechanical terms.     

A new stem craniate from the Ordovician of Morocco and the search for the sister group of the craniata

The investigation of the development of the trigeminal jaw adductor musculature in the turtle Chelydra serpentina documents the early aggregation of muscle rudiments around the innervating nerve branches, probably a consequence of inductive interaction. This may explain the early continuity of the intramandibularis with the intermandibularis muscle. Several aspects of muscle development differ in the turtle as compared to lizards. These differences highlight the fact that conjectures of homology, based on a static topographical correspondence of adult structures, cannot capture the dynamics of the developmental process. The intramandibularis muscle of turtles, comparable to that of crocodiles, represents a plesiomorphous structure which is not homologous to the intramandibularis muscle of lacertoid lizards, a derived feature of the Lacertoidea. A derived feature of the chelonian jaw adductor musculature is the posterodorsal expansion of the external adductor along a supraoccipital crest, developing according to a pattern of Haeckelian recapitulation. Muscle development serves to corroborate the concept of a monophyletic Eureptilia, including diapsids and synapsids, as opposed to the (paraphyletic) Anapsida. The impact of the differentiation of the external adductor into a pulley system on cranial kinesis is analysed in biomechanical terms.     

The trigeminal jaw adductors of primitive snakes and their homologies with the lacertilian jaw adductors

The trigeminal jaw adductor musculature of anilioid snakes is analysed. The group is characterised by primitive characters, viz. the presence of an extensive bodenaponeurosis and of a quadrate aponeurosis. A temporal tendon gives rise to superficial (lb) fibres which are not observed in other snakes: this may be a primitive or a derived feature.
Jaw adductor muscles in snakes are usually subdivided following their relative position in an antero–posterior direction. Lacertilian jaw adductors are subdivided in a transverse plane. A detailed comparison of the anilioid and primitive lacertilian jaw adductors establishes correspondences (homologies) of parts in the transverse plane in both groups. These homologies are corroborated by innervational patterns.
Platynotan lizards are widely accepted as potential snake ancestors. A comparison of homologue jaw adductors shows different evolutionary trends to characterise platynotan lizards and snakes. Theoretically, these findings do not rule out primitive platynotan lizards as snake ancestors. On the basis of the structure of jaw adductors, snakes are to be derived from a primitive lacertilian pattern, be it platynotan or not.     

Cranial kinesis in lizards (Lacertilia): Origin,biomechanics, and evolution

Various methods of investigation of cranial kinesis are compared. The biomechanical model of the amphikinetic cranial mechanism of lizards developed by Frazzetta (1962) corresponds to the skulls of species with the dependent streptostyly. A new modification of this model is proposed for species with the independent streptostyly, which conforms to the results of experimental investigations of cranial kinesis in living lizards. The lacertilian amphikinesis has developed on the basis of the pleurokinesis inherited from fish ancestors of tetrapods. Movable connections of the maxillo-buccal segments with the axial skull persisted in the amphikinetic skull and were completed by the transversal flexible connections in the dermatocranial roof and loose connections of dermatocranium with the braincase. The development of new movable intracranial connections could have been preceded by transformations in the jaw musculature (formation of the pterygoideus muscle and inclined position of the external jaw adductor), which caused longitudinal jaw movements. Development of new movable connections within the skull was triggered by paedomorphosis processes. In various lacertilian groups, the cranial kinesis was improved by the development of various forms of streptostyly and flexipalatality.     

The structure of the skull and jaw adductor musculature in the Gekkota, with comments on the phylogenetic relationships of the Xantusiidae (Reptilia: Lacertilia)

The skull and trigeminal jaw adductor musculature of the lizard families Gekkonidae, Pygopodidae and Xantusiidae are described. The external jaw adductor shows a different structure in the Gekkonidae and Pygopodidae than is observed in other lizards, approached only by the Xantusiidae and Feyliniidae. Paedomorphosis seems to be involved in the differentiation of the jaw adductor musculature in the Gekkonidae. The Gekkonidae and Pygopodidae may be hypothesized to form a monophyletic group, the Gekkota, on the basis of numerous synapomorphies. Within the Gekkota, the Pygopodidae are the sister-group of the Gekkonidae and retain some plesiomorphous features which are absent in the latter. The Xantusiidae share few synapomorphies with the Gekkota on the one hand, and some with scincomorph lizards on the other, especially with the Lacertidae.     

Archosaur adductor chamber evolution: integration of musculoskeletal and topological criteria in jaw muscle homology

The homologies of jaw muscles among archosaurs and other sauropsids have been unclear, confounding interpretation of adductor chamber morphology and evolution. Relevant topological patterns of muscles, nerves, and blood vessels were compared across a large sample of extant archosaurs (birds and crocodylians) and outgroups (e.g., lepidosaurs and turtles) to test the utility of positional criteria, such as the relative position of the trigeminal divisions, as predictors of jaw muscle homology. Anatomical structures were visualized using dissection, sectioning, computed tomography (CT), and vascular injection. Data gathered provide a new and robust view of jaw muscle homology and introduce the first synthesized nomenclature of sauropsid musculature using multiple lines of evidence. Despite the great divergences in cephalic morphology among birds, crocodylians, and outgroups, several key sensory nerves (e.g., n. anguli oris, n. supraorbitalis, n. caudalis) and arteries proved useful for muscle identification, and vice versa. Extant crocodylians exhibit an apomorphic neuromuscular pattern counter to the trigeminal topological paradigm: the maxillary nerve runs medial, rather than lateral to M. pseudotemporalis superficialis. Alternative hypotheses of homology necessitate less parsimonious interpretations of changes in topology. Sensory branches to the rictus, external acoustic meatus, supraorbital region, and other cephalic regions suggest conservative dermatomes among reptiles. Different avian clades exhibit shifts in some muscle positions, but maintain the plesiomorphic, diapsid soft-tissue topological pattern. Positional data suggest M. intramandibularis is merely the distal portion of M. pseudotemporalis separated by an intramuscular fibrocartilaginous sesamoid. These adductor chamber patterns indicate multiple topological criteria are necessary for interpretations of soft-tissue homology and warrant further investigation into character congruence and developmental connectivity.     

Functional basis for sexual differences in bite force in the lizard Anolis carolinensis

In many species of lizards, males attain greater body size and have larger heads than female lizards of the same size. Often, the dimorphism in head size is paralleled by a dimorphism in bite force. However, the underlying functional morphological basis for the dimorphism in bite force remains unclear. Here, we test whether males are larger, and have larger heads and bite forces than females for a given body size in a large sample of Anolis carolinensis . Next, we test if overall head shape differs between the sexes, or if instead specific aspects of skull shape can explain differences in bite force. Our results show that A. carolinensis is indeed dimorphic in body and head size and that males bite harder than females. Geometric morphometric analyses show distinct differences in skull shape between males and females, principally reflecting an enlargement of the jaw adductor muscle chamber. Jaw adductor muscle mass data confirm this result and show that males have larger jaw adductors (but not jaw openers) for a given body and head size. Thus, the observed dimorphism in bite force in A. carolinensis is not merely the result of an increase in head size, but involves distinct morphological changes in skull structure and the associated jaw adductor musculature.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 111–119.     

The development of the trigeminal jaw adductor musculature and associated skull elements in the lizard Podarcis sicula

The ontogenetic development of the jaw adductor musculature in Podarcis sicula (Raftnesque) is described in detail and related to patterns of ossification in associated skull elements. It was found that the coronoid bone ossifies in continuity with the developing bodenaponeurosis, whereas the elements of the upper temporal arcade ossify prior to (and seemingly independently of) the realization of an attachment of the developing jaw adductors.
Various aspects of the development of the jaw adductors are discussed in the light of theoretical claims concerning conflicting models of ontogeny, as is the role of ontogenetic studies in the determination of character polarity. It is concluded that the present study supports the model of epigenetic development, and that ontogeny alone is insufficient for the assessment of character polarity. Instead, ontogeny must be interpreted in the context of phylogenetic hypotheses.     

Comments on the evolution of the jaw adductor musculature of snakes

The aim of this study is to provide a general view of the adductor musculature of the alethinophidian snakes. The aponeurotic system present in anilioid snakes is here described as being also present in colubroid and booid snakes. Although modified in various groups, this aponeurotic system retains the same topographical pattern in the anilioids, booids and colubroids, and is thus hypothesized to be homologous. An analysis of the aponeurotic system and related muscular bundles within the alethinophidian snakes is given. A new terminology is proposed for the jaw adductor muscles where the muscles levator anguli oris and adductor mandibulae externus superficialis (proper) of snakes (sensu Lakjer, 1926; Haas, 1962) retain these names even if this fails to reflect the presumed homologies with the bundles of the same name in lizards (see Rieppel, 1988b); the fibres originating from the temporal tendon in the Anilioidea, and presumed to form a bundle of composite nature (Rieppel, 1980b), are named the M. adductor mandibulae externus temporalis (lost by the Macrostomata); the M. adductor mandibulae externus medialis is a composite muscle in the Anilioidea (Rieppel, 1980b) which give rise to two different muscles in the ‘booids’, the M. adductor mandibulae externus medialis, pars anterior and the M. adductor mandibulae externus profundus, the former being secondarily lost by the Caenophidia which retains only fibres homologues of the 3b and 3c heads of the profundus layer of lizards; the so-called M. adductor mandibular externus profundus of snakes (sensu Lackjer, 1926; Haas, 1962) is also a composite muscle in the Anilioidea (Rieppel, 1980b), in the alethinophidians it is essentially made of fibres homologous with the posterior pinnate part of the medialis layer of lizards, and is here named the M. adductor mandibulae externus medialis, pars posterior. As a result from this analysis it follows that: (1) the Macrostomata are characterized by the downward extension of the fibres forming the M. adductor mandibulae externus medialis, pars anterior and the loss of the M. adductor mandibulae externus temporalis: (2) the Xenopeltidae are set apart from the remaining macrostomatan snakes by the retention of the M. levator anguli oris and of a well developed lateral sheet of the quadrate aponeurosis; (3) the ‘booids’ form a monophyletic group comprising only the Boidae and Bolyeriidae (with the exclusion of the Xenopeltidae and Tropidophiidae) which is characterized by a differentiated M. adductor mandibulae externus medialis, pars anterior inserting on the lateral surface of the compound bone via its own aponeurosis; (4) the Tropidophiidae are set apart from all other snakes by the peculiar course of their lateral head vein; however, they belong to the Caenophidia as they show a facial carotid artery which passes dorsally to the mandibular and maxillary branches of the trigeminus; (5) a possible additional character in favour of an Acrochordoidea + Colubroidea monophyletic unit may be given by the pattern of innervation of the jaw adductor muscles in these two taxa; (6) a new interpretation of the compressor glandulae muscular complex of Atractaspis resulted in a morphologically similar pattern to that of the viperids; the phylogenetic implications of such similarity are discussed in detail.     

Tooth size and arcadal length correlates in man

Intra-arcadal mesiodistal and buccolingual tooth size correlations were evaluated in a sample of 125 caucasoids with ideal occlusion. Dental dimensions were corrected for arcade mength (as a measure of jaw size) by a series of regression analyses of each mesiodistal dimension on the sum of the mesiodistal dimensions within each arcade. Regression coefficients of tooth dimension on arcade length were calculated to gain an insight into the dimensional sensitivity of individual teeth to arcade length variation. The data presented here suggest a strong association between arcadal length (jaw size) dependence, and the dimensional stability of individual teeth. When corrected for arcade length, a definite pattern of tooth size correlation emerges: postcanine maxillary and mandibular teeth are negatively correlated to the anterior teeth and are positively correlated to one another. The hypothesis is developed that anterior and postcanine teeth should be viewed as two separate and negatively size-correlated units, beyond the boundaries of the four morphological tooth classes. Recognition of this basic dichotomous size arrangement within each jaw allows for a reassessment of some of the problems associated with hominid dental evolution.     

Cranial musculature of South American Gekkonidae

The cranial myology of 13 South American geckos was compared and analyzed for taxonomic significance. The general pattern emerges that muscles in this group do not vary substantially from those of other lizards, except that the geckonids have fewer muscle layers. The former condition is particularly important with respect to the adductor musculature of the jaw, given its fundamental role in the chewing process. A great variety of lizards, both those with the geckonid bauplan and those with other morphologies exhibit similar basic structures in the jaw adductor muscles, despite significant differences in diet. There appears to be no direct correlation between diet and the morphology of head musculature of lizards. It is hypothesized that differences between and within bauplans can be ascribed to phylogenetic factors rather than to functional characteristics such as diet and life-styles. Twenty characteristics reflecting minor variations in the Gekkotan bauplan were selected for comparison in performing a cladistic analysis rooted on the sphaerodactylinid geckoes Coleodactylus amazonicus and C. septentrionalis. Groupings of muscular characteristics resulting from this analysis lead to different interpretations of taxonomic relationships from those derived from previous studies on the taxa examined. © 1996 Wiley-Liss, Inc.     

Sexual Selection and Dynamics of Jaw Muscle in Tupinambis Lizards

Sexual dimorphism patterns provide an opportunity to increase our understanding of trait evolution. Because selective forces may vary throughout the reproductive period, measuring dimorphism seasonally may be an interesting approach. An increased male head size may be important in intersexual and intrasexual interactions. In Tupinambis lizards, a big head is attributed in part to a large adductor muscle mass. Competition for mating can differ in species with different sex ratio and different degrees of sexual size dimorphism. We examined sexual differences in mass of the pterygoideus muscle, its temporal variation throughout the reproductive period and the relationship between muscle and reproductive condition in Tupinambis merianae and T. rufescens. We characterized sexual size dimorphism and sex ratio in both species. Mature males had larger jaw muscles than mature females in both species, mainly during the reproductive season. The dimorphism in jaw muscle was due to an increase in muscle mass in sexually active males. Seasonal increases in muscle mass and variation between immature and mature individuals suggest that the jaw muscle might be a secondary sexual character. We propose that the pterygoideus muscle may act as a signal of reproductive condition of males because it is associated with testis size and sperm presence. The patterns of sexual dimorphism in jaw muscle in both species were similar; however, the comparison shows how sexual characters remain dimorphic in different competition contexts and in species with different degrees of body size dimorphism. Our results suggest that jaw muscle as sexual character could be influenced by inter- and intrasexual selective pressures.     

More than 1000 ultraconserved elements provide evidence that turtles are the sister group of archosaurs

We present the first genomic-scale analysis addressing the phylogenetic position of turtles, using over 1000 loci from representatives of all major reptile lineages including tuatara. Previously, studies of morphological traits positioned turtles either at the base of the reptile tree or with lizards, snakes and tuatara (lepidosaurs), whereas molecular analyses typically allied turtles with crocodiles and birds (archosaurs). A recent analysis of shared microRNA families found that turtles are more closely related to lepidosaurs. To test this hypothesis with data from many single-copy nuclear loci dispersed throughout the genome, we used sequence capture, high-throughput sequencing and published genomes to obtain sequences from 1145 ultraconserved elements (UCEs) and their variable flanking DNA. The resulting phylogeny provides overwhelming support for the hypothesis that turtles evolved from a common ancestor of birds and crocodilians, rejecting the hypothesized relationship between turtles and lepidosaurs.     

Anatomy of the feeding apparatus of the lemon shark,Negaprion brevirostris

The anatomy of the feeding apparatus of the lemon shark, Negaprion brevirostris, is investigated by gross dissection, computer axial tomography, and histological staining. The muscles and ligaments of the head associated with feeding are described. The upper and lower jaws are suspended by the hyoid arch, which in turn is braced against the chondrocranium by a complex series of ligaments. In addition, various muscles and the integument contribute to the suspension and stability of the jaws. The dual jaw joint is comprised of lateral and medial quadratomandibular joints that resist lateral movement of the upper and lower jaws on one another. This is important during feeding involving vigorous head shaking. An elastic ethmoplatine ligament that unites the anterior portion of the upper jaw to the neurocranium is involved with upper jaw retraction. The quadratomandibularis muscle is divided into four divisions with a bipinnate fiber arrangement of the two large superficial divisions. This arrangement would permit a relatively greater force per unit volume and reduce muscle bulging of the jaw adductor muscle in the spatially confined cheek region. Regions of relatively diffuse integumental ligaments overlying the adductor mandibulae complex and the levator palatoquadrati muscle, interspersed with localized regions of longer tendonlike attachments between the skin and the underlying muscle, permit greater musculoskeletal movement relative to the skin. The nomenclature of the hypobranchial muscles is discussed. In this shark they are comprised of the unsegmented coracomandibularis and coracohyoideus, and the segmented coracoarcualis. © 1995 Wiley-Liss, Inc.     

Evolution and mechanics of long jaws in butterflyfishes (family Chaetodontidae)

We analyzed the functional morphology and evolution of the long jaws found in several butterflyfishes. We used a conservative reanalysis of an existing morphological dataset to generate a phylogeny that guided our selection of seven short- and long-jawed taxa in which to investigate the functional anatomy of the head and jaws: Chaetodon xanthurus, Prognathodes falcifer (formerly Chaetodon falcifer), Chelmon rostratus, Heniochus acuminatus, Johnrandallia nigrirostris, Forcipiger flavissimus, and F. longirostris. We used manipulations of fresh, preserved, and cleared and stained specimens to develop mechanical diagrams of how the jaws might be protruded or depressed. Species differed based on the number of joints within the suspensorium. We used high-speed video analysis of five of the seven species (C. xanthurus, Chel. rostratus, H. acuminatus, F. flavissimus, and F. longirostris) to test our predictions based on the mechanical diagrams: two suspensorial joints should facilitate purely anteriorly directed protrusion of the lower jaw, one joint should allow less anterior protrusion and result in more depression of the lower jaw, and no joints in the suspensorium should constrain the lower jaw to simple ventral rotation around the jaw joint, as seen in generalized perciform fishes. We found that the longest-jawed species, F. longirostris, was able to protrude its jaws in a predominantly anterior direction and further than any other species. This was achieved with little input from cranial elevation, the principal input for other known lower jaw protruders, and is hypothesized to be facilitated by separate modifications to the sternohyoideus mechanism and to the adductor arcus palatini muscle. In F. longirostris the adductor arcus palatini muscle has fibers oriented anteroposteriorly rather than medial-laterally, as seen in most other perciforms and in the other butterflyfish studied. These fibers are oriented such that they could rotate the ventral portion of the quadrate anteriorly, thus projecting the lower jaw anteriorly. The intermediate species lack modification of the adductor arcus palatini and do not protrude their jaws as far (in the case of F. flavissimus) or in a purely anterior fashion (in the case of Chel. rostratus). The short-jawed species both exhibit only ventral rotation of the lower jaw, despite the fact that H. acuminatus is closely related to Forcipiger.     

Control of jaw movements in two species of macropodines (Macropus eugenii and Macropus rufus)

The masticatory motor patterns of three tammar wallabies and two red kangaroos were determined by analyzing the pattern of electromyographic (EMG) activity of the jaw adductors and correlating it with lower jaw movements, as recorded by digital video and videoradiography. Transverse jaw movements were limited by the width of the upper incisal arcade. Molars engaged in food breakdown during two distinct occlusal phases characterized by abrupt changes in the direction of working-side hemimandible movement. Separate orthal (Phase I) and transverse (Phase II) trajectories were observed. The working-side lower jaw initially was drawn laterally by the balancing-side medial pterygoid and then orthally by overlapping activity in the balancing- and working-side temporalis and the balancing-side superficial masseter and medial pterygoid. Transverse movement occurred principally via the working-side medial pterygoid and superficial masseter. This pattern contrasted to that of placental herbivores, which are known to break down food when they move the working-side lower jaw transversely along a relatively longer linear path without changing direction during the power stroke. The placental trajectory results from overlapping activity in the working- and balancing-side adductor muscles, suggesting that macropods and placental herbivores have modified the primitive masticatory motor pattern in different ways.     

Jaw movement kinematics and jaw muscle (EMG) activity during drinking in the pigeon (Columba livia)

Summary Movements of the maxilla and mandible were recorded during drinking in the head-fixed pigeon and correlated with electromyographic activity in representative jaw muscle groups. During drinking, each jaw exhibits opening and closing movements along both the dorso-ventral and rostro-caudal axes which may be linked with or independent of each other. All subjects showed small but systematic increases in cycle duration over the course of individual drinking bouts. Cyclic jaw movements during drinking were correlated with nearly synchronous activity in the protractor (levator) of the upper jaw and in several jaw closer muscles, as well as with alternating activity in tongue protractor and retractor muscles. No EMG activity was ever recorded in the lower jaw opener muscle, suggesting that lower jaw opening in this preparation is produced, indirectly, by the contraction of other muscles. The results clarify the contribution of the individual jaws to the generation of gape variations during drinking in this species.Abbreviations AMEM adductor mandibulae externus muscle - DM depressor mandibulae muscle - EMG electromyographic - GENIO geniohyoideus muscle - LB lower beak - LED light-emitting diode - PQP protractor quadrati et pterygoidei muscle - PVL pterygoideus ventralis muscle, pars lateralis - SeH/StH serpihyoideus or stylohyoideus muscle - UB upper beak     

Feeding mechanics in Triassic stem-group sauropterygians: the anatomy of a successful invasion of Mesozoic seas

The jaw adductor musculature in Triassic stem-group sauropterygians is reconstructed on the basis of a paradigmatic model of muscle architecture (functional equivalence of sarcomeres) and using invariant traits of the anatomy of the trigeminal jaw adductor muscles in extant reptiles. The reconstructed jaw adductor musculature predicts trophic specializations in stem-group sauropterygians. Suction feeding is a component in prey capture for some benthic feeding, as well as for some pelagic feeding taxa. The differentiation of 'pincer' jaws is correlated with the potential for rapid, snapping bites. There is some evidence for habitat partitioning among Triassic stem-group sauropterygians with respect to trophic specialization. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 135 , 33–63.     

Control of jaw movements in two species of macropodines (Macropus eugenii and Macropus rufus).

The masticatory motor patterns of three tammar wallabies and two red kangaroos were determined by analyzing the pattern of electromyographic (EMG) activity of the jaw adductors and correlating it with lower jaw movements, as recorded by digital video and videoradiography. Transverse jaw movements were limited by the width of the upper incisal arcade. Molars engaged in food breakdown during two distinct occlusal phases characterized by abrupt changes in the direction of working-side hemimandible movement. Separate orthal (Phase I) and transverse (Phase II) trajectories were observed. The working-side lower jaw initially was drawn laterally by the balancing-side medial pterygoid and then orthally by overlapping activity in the balancing- and working-side temporalis and the balancing-side superficial masseter and medial pterygoid. Transverse movement occurred principally via the working-side medial pterygoid and superficial masseter. This pattern contrasted to that of placental herbivores, which are known to break down food when they move the working-side lower jaw transversely along a relatively longer linear path without changing direction during the power stroke. The placental trajectory results from overlapping activity in the working- and balancing-side adductor muscles, suggesting that macropods and placental herbivores have modified the primitive masticatory motor pattern in different ways.