Comparative myology of the mandibular and hyoid arches of sharks of the order hexanchiformes and their bearing on its monophyly and phylogenetic relationships (Chondrichthyes: Elasmobranchii)

The order Hexanchiformes currently comprises two families, Chlamydoselachidae (frilled sharks) and Hexanchidae (six‐ and seven‐gill sharks), but its monophyly and relationships with other elasmobranchs are still discussed. Previous studies of hexanchiforms addressing these issues were based mainly on external morphology, teeth, skeletal features, and molecular data, whereas the employment of characters derived from variations in muscles has not been significantly explored. Dissections of four species of Hexanchiformes (including Chlamydoselachus anguineus) are reported here describing the mandibular (musculus adductor mandibulae dorsalis, m. adductor mandibulae ventralis, m. levator labii superioris, m. intermandibularis, and m. constrictor dorsalis) and hyoidean (m. constrictor hyoideus dorsalis and ventralis) arch muscles. Our results provide new data concerning the relationships of hexanchiforms to other elasmobranchs. The m. adductor mandibulae superficialis is described and illustrated in C. anguineus, contradicting previous accounts in which is was considered absent. The anteroposterior orientation of the m. adductor mandibulae superficialis in Chlamydoselachus is similar to the pattern found in hexanchids, squaloids, and hypnosqualeans (including batoids), suggesting it was secondarily lost in Echinorhinus. This muscle therefore provides further support for the inclusion of the Chlamydoselachidae and Hexanchidae in the Squalomorphi, and not basal to all other elasmobranchs or nested within an all‐shark collective, as has been previously proposed. However, the m. adductor mandibulae superficialis originating at the jaw joint and with an aponeurotic insertion in hexanchids, squaliforms, and hypnosqualeans, may be a separate derived feature uniting these taxa. The insertion of the m. constrictor dorsalis is restricted to the postorbital articulation in hexanchids, whereas it extends farther anteriorly in C. anguineus. The insertion of the m. constrictor hyoideus dorsalis solely on the palatoquadrate is found exclusively in the Hexanchidae. We conclude that no specific pattern of mandibular or hyoid arch muscles support the monophyly of hexanchiforms (i.e., including Chlamydoselachus). J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

The complete mitochondrial genome sequence of the world's largest fish,the whale shark (Rhincodon typus), and its comparison with those of related shark species

The whale shark (Rhincodon typus) is the largest extant species of fish, belonging to the order Orectolobiformes. It is listed as a “vulnerable” species on the International Union for Conservation of Nature (IUCN)'s Red List of Threatened Species, which makes it an important species for conservation efforts. We report here the first complete sequence of the mitochondrial genome (mitogenome) of the whale shark obtained by next-generation sequencing methods. The assembled mitogenome is a 16,875 bp circle, comprising of 13 protein-coding genes, two rRNA genes, 22 tRNA genes and a control region. We also performed comparative analysis of the whale shark mitogenome to the available mitogenome sequences of 17 other shark species, four from the order Orectolobiformes, five from Lamniformes and eight from Carcharhiniformes. The nucleotide composition, number and arrangement of the genes in whale shark mitogenome are the same as found in the mitogenomes of the other members of the order Orectolobiformes and its closest orders Lamniformes and Carcharhiniformes, although the whale shark mitogenome had a slightly longer control region. The availability of mitogenome sequence of whale shark will aid studies of molecular systematics, biogeography, genetic differentiation, and conservation genetics in this species.     

Shark tales: a molecular species-level phylogeny of sharks (Selachimorpha, Chondrichthyes)

Sharks are a diverse and ecologically important group, including some of the ocean's largest predatory animals. Sharks are also commercially important, with many species suffering overexploitation and facing extinction. However, despite a long evolutionary history, commercial, and conservation importance, phylogenetic relationships within the sharks are poorly understood. To date, most studies have either focused on smaller clades within sharks, or sampled taxa sparsely across the group. A more detailed species-level phylogeny will offer further insights into shark taxonomy, provide a tool for comparative analyses, as well as facilitating phylogenetic estimates of conservation priorities. We used four mitochondrial and one nuclear gene to investigate the phylogenetic relationships of 229 species (all eight Orders and 31 families) of sharks, more than quadrupling the number of taxon sampled in any prior study. The resulting Bayesian phylogenetic hypothesis agrees with prior studies on the major relationships of the sharks phylogeny; however, on those relationships that have proven more controversial, it differs in several aspects from the most recent molecular studies. The phylogeny supports the division of sharks into two major groups, the Galeomorphii and Squalimorphii, rejecting the hypnosqualean hypothesis that places batoids within sharks. Within the squalimorphs the orders Hexanchiformes, Squatiniformes, Squaliformes, and Pristiophoriformes are broadly monophyletic, with minor exceptions apparently due to missing data. Similarly, within Galeomorphs, the orders Heterodontiformes, Lamniformes, Carcharhiniformes, and Orectolobiformes are broadly monophyletic, with a couple of species 'misplaced'. In contrast, many of the currently recognized shark families are not monophyletic according to our results. Our phylogeny offers some of the first clarification of the relationships among families of the order Squaliformes, a group that has thus far received relatively little phylogenetic attention. Our results suggest that the genus Echinorhinus is not a squaliform, but rather related to the saw sharks, a hypothesis that might be supported by both groups sharing 'spiny' snouts. In sum, our results offer the most detailed species-level phylogeny of sharks to date and a tool for comparative analyses.     

Development of the muscles associated with the mandibular and hyoid arches in the Siberian sturgeon,Acipenser baerii (Acipenseriformes: Acipenseridae)

The skeleton of the jaws and neurocranium of sturgeons (Acipenseridae) are connected only through the hyoid arch. This arrangement allows considerable protrusion and retraction of the jaws and is highly specialized among ray‐finned fishes (Actinopterygii). To better understand the unique morphology and the evolution of the jaw apparatus in Acipenseridae, we investigated the development of the muscles of the mandibular and hyoid arches of the Siberian sturgeon, Acipenser baerii. We used a combination of antibody staining and formalin‐induced fluorescence of tissues imaged with confocal microscopy and subsequent three‐dimensional reconstruction. These data were analyzed to address the identity of previously controversial and newly discovered muscle portions. Our results indicate that the anlagen of the muscles in A. baerii develop similarly to those of other actinopterygians, although they differ by not differentiating into distinct muscles. This is exemplified by the subpartitioning of the m. adductor mandibulae as well as the massive m. protractor hyomandibulae, for which we found a previously undescribed portion in each. The importance of paedomorphosis for the evolution of Acipenseriformes has been discussed before and our results indicate that the muscles of the mandibular and the hyoid may be another example for heterochronic evolution.     

Morphology and evolution of the jaw suspension in lamniform sharks

The morphology of the jaw suspension and jaw protrusion mechanism in lamniform sharks is described and mapped onto a cladogram to investigate how changes in jaw suspension and protrusion have evolved. This has revealed that several evolutionary modifications in the musculoskeletal apparatus of the jaws have taken place among lamniform sharks. Galeomorph sharks (Carcharhiniformes, Lamniformes, Orectolobiformes, and Heterodontiformes) have paired ethmopalatine ligaments connecting the ethmoid process of the upper jaw to the ethmoid region of the cranium. Basal lamniform sharks also acquired a novel single palatonasal ligament connecting the symphysis of the upper jaw to the cranium mid-ventral to the nasal capsule. Sharks in the family Lamnidae subsequently lost the original paired ethmopalatine ligament while retaining the novel palatonasal ligament. Thus, basal lamniform taxa (Mitsukurina owstoni, Carcharius taurus, Alopias vulpinnis) have increased ligamentous support of the lateral region of the upper jaw while derived species (Lamnidae) have lost this lateral support but gained anterior support. In previous studies the morphology of the jaw suspension has been shown to play a major role in the mechanism of upper jaw protrusion in elasmobranchs. The preorbitalis is the primary muscle effecting upper jaw protrusion in squalean (sister group to galeomorphs) and carcharhiniform (sister group to lamniforms) sharks. The preorbitalis originates from the quadratomandibularis muscle and inserts onto the nasal capsule in squalean and carcharhiniform sharks. Carcharhiniform sharks have evolved a subdivided preorbitalis muscle with the new division inserting near the ethmoid process of the palatoquadrate (upper jaw). Alopid sharks have also independently evolved a partially subdivided preorbitalis with the new division inserting at the base of the ethmoid process and surrounding connective tissue. Lamnid sharks have retained the two preorbitalis divisions but have modified both of the insertion points. The original ventral preorbitalis division now inserts onto the connective tissue surrounding the mid-region of the upper jaw, while the new dorsal preorbitalis division inserts onto the surrounding connective tissue and skin at a more posterior position on the upper jaw. The retractor muscle of the jaws, the levator hyomandibularis, has also been modified during the evolution of lamniform sharks. In most sharks, including basal lamniforms, the levator hyomandibularis inserts onto the hyomandibula and functions to retract the jaws after protrusion. In alopid and lamnid sharks the levator hyomandibularis inserts primarily onto the upper and lower jaws around the jaw joint and is a more direct route for retracting the jaws. Thus, there has been at least one instance of character loss (ethmopalatine ligament), acquisition (palatonasal ligament), subdivision (preorbitalis), and modification (ventral preorbitalis, dorsal preorbitalis, and levator hyomandibularis) in the ligaments and muscles associated with the jaw suspension and jaw protrusion mechanism in lamniform sharks. While derived lamniform sharks (Lamna nasus, Carcharodon carcharius, and Isurus oxyrinchus) lost the ancestral passive lateral support of the ethmoid articulation of the upper jaw, they simultaneously acquired muscular support by way of the levator hyomandibularis, which provides a dynamic mechanism for lateral support. The evolution of multiple divisions of preorbitalis insertions onto the palatoquadrate and modification of the levator hyomandibularis insertion directly onto the jaws provides an active mechanism for multiple protractions and retractions of the upper jaw, which is advantageous in those sharks that gouge or saw pieces from large oversized prey items.     

A new method for correcting a gummy smile

Myectomy and partial resection of the levator labii superioris promises a far better and direct approach to the problem of upper gum exposure during smiling. This procedure attempts to eliminate the cause of the deformity. The resected muscles are of the striated muscle group and are composed of single muscle fibers running the entire length of the trunk. Adequate resection of these muscles will thoroughly eliminate the regeneration of muscle, and thus a permanent correction is achieved. The resulting smile is very attractive and pleasing to both patient and surgeon.     

Molecular phylogeny of elasmobranchs inferred from mitochondrial and nuclear markers

The elasmobranchs (sharks, rays and skates) being the extant survivors of one of the earliest offshoots of the vertebrate evolutionary tree are good model organisms to study the primitive vertebrate conditions. They play a significant role in maintaining the ecological balance and have high economic value. Due to over-exploitation and illegal fishing worldwide, the elasmobranch stocks are being decimated at an alarming rate. Appropriate management measures are necessary for restoring depleted elasmobranch stocks. One approach for restoring stocks is implementation of conservation measures and these measures can be formulated effectively by knowing the evolutionary relationship among the elasmobranchs. In this study, a total of 30 species were chosen for molecular phylogeny studies using mitochondrial cytochrome c oxidase subunit I, 12S ribosomal RNA gene and nuclear Internal Transcribed Spacer 2. Among different genes, the combined dataset of COI and 12S rRNA resulted in a well resolved tree topology with significant bootstrap/posterior probabilities values. The results supported the reciprocal monophyly of sharks and batoids. Within Galeomorphii, Heterodontiformes (bullhead sharks) formed as a sister group to Lamniformes (mackerel sharks): Orectolobiformes (carpet sharks) and to Carcharhiniformes (ground sharks). Within batoids, the Myliobatiformes formed a monophyly group while Pristiformes (sawfishes) and Rhinobatiformes (guitar fishes) formed a sister group to all other batoids.     

A simplified method for smile enhancement: botulinum toxin injection for gummy smile

The authors studied and provided treatment to patients with the complaint of gummy smile. Between October of 2009 and January of 2011, 52 unaesthetic smiles were evaluated and treated with onabotulinum toxin A. Botulinum toxin injection was an effective treatment, with an average satisfaction level of 9.75 on a 10-point scale. The levator labii superioris alaeque nasi is the ideal muscle for injection, and lopsided smiles could be resolved also with onabotulinum toxin A asymmetric injection. Gummy smile treatment with onabotulinum toxin A into the levator labii superioris alaeque nasi muscle is an effective method, with minimum risk of complications and very high patient satisfaction.     

Anatomy of the feeding apparatus of the nurse shark,Ginglymostoma cirratum

The anatomy of the feeding apparatus of the nurse shark, Ginglymostoma cirratum, was investigated by gross dissection and computer axial tomography. The labial cartilages, jaws, jaw suspension, muscles, and ligaments of the head are described. Palatoquadrate cartilages articulate with the chondrocranium caudally by short, laterally projecting hyomandibulae and rostrally by ethmoorbital articulations. Short orbital processes of the palatoquadrates are joined to the ethmoid region of the chondrocranium by short, thin ethmopalatine ligaments. In addition, various ligaments, muscles, and the integument contribute to the suspension of the jaws. When the mouth is closed and the palatoquadrate retracted, the palatine process of the palatoquadrate is braced against the ventral surface of the nasal capsule and the ascending process of the palatoquadrate is in contact with the rostrodorsal end of the suborbital shelf. When the mandible is depressed and the palatoquadrate protrudes slightly rostroventrally, the palatoquadrate moves away from the chondrocranium. A dual articulation of the quadratomandibular joint restricts lateral movement between the mandible and the palatoquadrate. The vertically oriented preorbitalis muscle spans the gape and is hypothesized to contribute to the generation of powerful crushing forces for its hard prey. The attachment of the preorbitalis to the prominent labial cartilages is also hypothesized to assist in the retraction of the labial cartilages during jaw closure. Separate levator palatoquadrati and spiracularis muscles, which are longitudinally oriented and attach the chondrocranium to the palatoquadrate, are hypothesized to assist in the retraction of the palatoquadrate during the recovery phase of feeding kinematics. Morphological specializations for suction feeding that contribute to large subambient suction pressures include hypertrophied coracohyoideus and coracobranchiales muscles to depress the hyoid and branchial arches, a small oral aperture with well‐developed labial cartilages that occlude the gape laterally, and small teeth. J. Morphol. 241:33–60, 1999. © 1999 Wiley‐Liss, Inc.     

The hyomandibulae of rhizodontids (Sarcopterygii, stem-tetrapoda)

Despite its important role in the study of the evolution of tetrapods, the hyomandibular bone (the homologue of the stapes in crown-group tetrapods) is known for only a few of the fish-like members of the tetrapod stem-group. The best-known example, that of the tristichopterid Eusthenopteron, has been used as an exemplar of fish-like stem-tetrapod hyomandibula morphology, but in truth the conditions at the base of the tetrapod radiation remain obscure. We report, here, four hyomandibulae, from three separate localities, which are referable to the Rhizodontida, the most basal clade of stem-tetrapods. These specimens share a number of characteristics, and are appreciably different from the small number of hyomandibulae reported for other fish-like stem-tetrapods. While it is unclear if these characteristics represent synapomorphies or symplesiomorphies, they highlight the morphological diversity of hyomandibulae within the early evolution of the tetrapod total-group. Well-preserved muscle scarring on some of these hyomandibulae permit more robust inferences of hyoid arch musculature in stem-tetrapods.     

Homologies of the branchial arch muscles in Zacco platypus (Teleostei: Cypriniformes: Cyprinidae): Evidence from innervation pattern

Homologies of the branchial arch muscles in the cyprinid Zacco platypus are assessed based on their innervation. Muscles serving the first gill arch are innervated by branches of the glossopharyngeal (IX) nerve and those serving other arches by the vagal (X) nerve. Absence of the levator posterior is confirmed. Five pairs of muscles originating from the cranium and inserted onto the specialized 5th ceratobranchial, all unique to cyprinids, are innervated by the 4th branchial trunks of X, indicating that all pairs are derivatives of the sphincter oesophagi, involving reorganization from intrinsic to extrinsic elements. Homologies of some ventral branchial muscles are also discussed and the criteria for homology improved by clarifying the innervation pattern. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

On the muscles of expression in the geladaTheropithecus gelada (Rüppell) (primates,Cercopithecidae)

Musculature innervated by the N. facialis inTheropithecus gelada (Rüppell) is patterned on broad lines in agreement with related genera of catarrhine monkeys, but presents some specializations and divergences in detail. Noteworthy is the extension to the labial margins superficially of the combined levator labii superioris and zygomaticus in the upper and the pars mandibularis of trachelo-platysma in the lower lip. A specialization of the medial fibres of levator labii superioris forms a sling-like structure within the upper lip and serves to implement the lip-flip gesture characteristic of the genus. Its antagonist is the orbicularis oris. Special features of all other facialis muscles are considered.Abbreviations ABD Anterior belly of digastricus - AE Arteria facialis - ALS Arteria labialis superior - ANL Arteria lateralis nasi - AP Auricularis posterior - APA Arteria auricularis posterior - AS Arteria auricularis superior - A.Se. Arteria septi nasi - A.Sy. Arteria symphysialis - BP Buccal pouch - LAO Depressor anguli oris - FTA Fronto-temporo-auricularis - GLI Glandulae labialis inferiores - GLS Glandulae labialis superioris - LAO Levator anguli oris - LG Artery to labial glands - LLAN Levator labii superioris alaeque nasi - LLS Levator labii superioris - M Masseter - MM Musculus mentalis - NP Notoplatysma - O Occipitalis - OO Orbicularis oris - O.Oc Orbicularis oculi - P Procerus - SH Sterno-hyoideus - TP Trachelo-platysma - VL Vena labialis communis - VP Venous plexus of dorsum nasi - ZM Zygomaticus minor - Zy Zygomaticus     

Testing morphologically based phylogenetic theories within the cartilaginous fishes with molecular data, with special reference to the catshark family (Chondrichthyes; Scyliorhinidae) and the interrelationships within them

A molecular phylogenetic investigation was conducted to examine phylogenetic relationships between various members of the catsharks (Chondrichthyes; Carcharhiniformes; Scyliorhinidae), and is the largest chondrichthyan data set yet analysed, consisting of nearly 130,000 nucleotides. Three mitochondrial DNA genes were used to construct the phylogenies, cytochrome b, NADH-2, and NADH-4, with 41 sequences from 18 taxa being novel. These sequences were either used separately or combined into a single data set, and phylogenies were constructed using various methods, however, only the Bayesian inference tree derived from the cytochrome b data set was resolved sufficiently for phylogenetic inferences to be made. Interestingly, the family Scyliorhinidae was not supported by the results and was found to be paraphyletic. The Scyliorhininae and Pentanchinae were supported, whereas the Pentanchini clade was present, but not well supported. The Halaelurini hypothesis was supported with Holohalaelurus identified as the basal genus of that clade, and Haploblepharus edwardsii identified as the basal taxon for that genus. Elsewhere within the Chondrichthyes, the Carcharhiniformes and the Lamniformes were found to be monophyletic, and the Heterodontiformes was placed within the Squalimorphs. The placement of the skates and rays in these analyses support the Batoidea as being sister to the Elasmobranchii.     

Neuromuscular spindles of the extrinsic ocular musculature of the moufflon

Proprioceptive innervation of moufflon extrinsic ocular musculature and m. levator palpebrae superioris was studied. Muscle spindles and Golgi tendon organs were found. The first ones are usually between 1st and 2nd order muscle fascicles. The muscle spindles are highly represented in the extrinsic ocular muscles, but less numerous in m. levator palpebrae superioris. Their number varies according to muscles and individuals. In the same subject, also the ratio between the number of the muscle spindles found in m. rectus dorsalis and that of m. levator palpebrae superioris was examined. Besides, the histological structure of the intrafusal fascicles was investigated. Particular attention was devoted to the nerve supply of the muscle spindle. By means of impregnating methods, sensory and motor endings were identified. Primary and secondary sensory endings only in a few cases showed their usual pattern: motor fibres can end in form of plates or trails. Golgi tendon organs were observed between the tendon and the muscular tissue and are always less numerous.     

Predilection sites of Trichinella spiralis larvae in naturally infected horses.

A total of 120 muscle tissues from three horses naturally infected with Trichinella spiralis were examined. The head was the most infected site. In particular, the muscles harbouring the highest number of larvae were: musculus buccinator (12, 411 and 1183 larvae g-1), the tongue (11, 615 and 1749 larvae g-1), m. levator labii maxillaris (17,582 and 1676 larvae g-1), and the masseter (4.9, 289 and 821 larvae g-1). Compared with the diaphragm, the number of larvae per gram was from 3.5 to 6.8 times higher in the tongue, from 3.5 to 6.5 higher in m. levator labii maxillaris, and from 2.5 to 4.6 higher in m. buccinator. Of the examined muscles, the diaphragm had from the 6th to the 15th highest level of infection (3.1, 166 and 256 larvae g-1). Published data from experimentally infected horses confirm these results, suggesting that efforts to detect predilection sites should focus on the head muscles.     

Dynamic rhinoplasty for the plunging nasal tip: functional unity of the inferior third of the nose

To achieve permanent results for the correction of a drooping nasal tip, it is important to understand the mechanism responsible for the caudal rotation of the tip when a person speaks or smiles. This mechanism can be considered to depend on a "functional unity" formed by three components: (1) the cartilaginous framework (alar cartilages and accessories acting as a single structure); (2) muscular motors (m. levator labii superioris alaeque nasi and depressor septi nasi); and (3) gliding areas (apertura piriformis, the valvular mechanism between the upper lateral cartilages and alar cartilages, the lax tissue of the nasal dorsum, and the membranous septum). We describe a new anatomical and functional concept responsible for the plunging of the nasal tip. When a person smiles, the functional unit is activated by a combination of two forces acting simultaneously in opposite directions that rotate the tip caudally and elevate the nasal base. The levator moves the alar base upward and the depressor pulls the tip caudally. To correct the drooping tip, the transcartilaginous incision is extended laterally, and the lateral portion of the alar arch is dissected free from the skin and the mucosa, thus exposing the accessory cartilages. The arch is then severed at the level of the accessories to allow the cephalad rotation of the domes. The muscle insertions are dissected free from the accessories and a section of the muscle and, if necessary, the accessory cartilages, is removed. From January of 1991 onward, 312 patients have had this ancillary procedure performed in addition to the basic rhinoplasty technique.     

Evolution of muscle activity patterns driving motions of the jaw and hyoid during chewing in Gnathostomes

Although chewing has been suggested to be a basal gnathostome trait retained in most major vertebrate lineages, it has not been studied broadly and comparatively across vertebrates. To redress this imbalance, we recorded EMG from muscles powering anteroposterior movement of the hyoid, and dorsoventral movement of the mandibular jaw during chewing. We compared muscle activity patterns (MAP) during chewing in jawed vertebrate taxa belonging to unrelated groups of basal bony fishes and artiodactyl mammals. Our aim was to outline the evolution of coordination in MAP. Comparisons of activity in muscles of the jaw and hyoid that power chewing in closely related artiodactyls using cross-correlation analyses identified reorganizations of jaw and hyoid MAP between herbivores and omnivores. EMG data from basal bony fishes revealed a tighter coordination of jaw and hyoid MAP during chewing than seen in artiodactyls. Across this broad phylogenetic range, there have been major structural reorganizations, including a reduction of the bony hyoid suspension, which is robust in fishes, to the acquisition in a mammalian ancestor of a muscle sling suspending the hyoid. These changes appear to be reflected in a shift in chewing MAP that occurred in an unidentified anamniote stem-lineage. This shift matches observations that, when compared with fishes, the pattern of hyoid motion in tetrapods is reversed and also time-shifted relative to the pattern of jaw movement.     

Gill‐arch musculature of the quillback carpiodes cyprinus (cypriniformes: catostomidae) with a comparison to cyprinids

Although the gill‐arch osteology of Cypriniformes has been well studied, comparable works on gill‐arch musculature are scarce. The focus of previous studies has been on Cyprinidae while other families have received little or no attention. Consequently, generalizations for Cypriniformes have been made from the musculature of cyprinid gill‐arches. This study describes the gill‐arch musculature of a catostomid, the quillback Carpiodes cyprinus, and demonstrates that there are striking differences in the overall gill‐arch musculature of catostomids in comparison to cyprinids, especially in the dorsal gill‐arch region. Of the 23 muscles found in the dorsal gill‐arch region of cyprinids, only 13 were present in C. cyprinus. Muscles that are absent include adductores 1–5, levator internus 4, levator ceratobranchialis 5 accessorius, retractor ceratobranchialis 5 externus, retractor ceratobranchialis 5 internus, and the retractor ceratobranchialis 5 transversus. In the ventral gill‐arch region, the rectus communis is absent. The derived scrolling shape of the dorsal gill‐arch skeleton associated with food processing is likely related to the change in musculature. J. Morphol., 2012. © 2012 Wiley Periodicals, Inc.     

Hyoid and pharyngeal arch function during ventilation and feeding in elasmobranchs: Conservation and modification in function

The hypothesis that the mandibular and hyoid arches evolved from anterior pharyngeal arches to increase ventilation performance and subsequently became adapted for feeding is widely accepted. As jaws evolved, the morphology of the hyoid arch changed notably from that of a pharyngeal arch. Furthermore, hyoid arch morphology varies considerably among elasmobranch taxa and has been shown to be related to feeding style. The goal of this study is to determine whether the function (direction of movement or change in cavity cross‐section) of the hyoid arch is altered from that of the pharyngeal arch, and whether function is altered between ventilation, the basal behavior, and feeding, the derived behavior. Similar effects and associations of the pharyngeal arches by orientation to feeding or ventilation are also investigated. The kinematics of the hyoid and second pharyngeal arch during ventilation and feeding are quantified using sonomicrometry and hyomandibular angle measured in five shark and one skate species representing widely divergent hyomandibular morphologies. Hyoid and pharyngeal cavity width follows the same pattern of movement during ventilation; therefore the hyoid arch retains the ancestral function of the pharyngeal arches. The orientation of the hyomandibular cartilage appears to influence the pattern of arch movement during ventilation: anterior directed elements decrease in cavity width; laterally directed elements increase in cavity width; while posterior directed elements increase in cavity width or do not change; while cavity depth increases in all species. Hyoid and pharyngeal cavity width movement differs among the species during feeding and also appears to be related to hyoid arch orientation as well as feeding style. There appears to be a division between those species with hyomandibular angles less than 110° from those that are greater between feeding mode and hyoid cavity width movement. Primarily suction feeding species decrease hyoid cavity width whereas primarily bite feeding species increase hyoid cavity width during feeding while all species increase hyoid cavity depth.