Hyoid Bar Research Articles

A saw-toothed eel †Serrivomer glehni sp. nov. from the Miocene of Sakhalin Island, north-western Pacific

ABSTRACT †Serrivomer glehni, a new species of the Neogene saw-toothed eel (Anguilliformes, Serrivomeridae) is described based on the 55 specimens from the Middle–Upper Miocene Kurasi Formation of Sakhalin Island, Russia. This is the third record of fossil saw-toothed eels since the recovery of partial skeletons from the Miocene of Japan and Italy. The new species demonstrates close similarity to the extant Serrivomer species in overall body form and shape of separate bones and their complexes. The new species is characterized by the “longidentatus” type of vomerine dentition and by a single branchiostegal ray protruding beyond the hyoid bar, which suggests a close affinity to the modern S. sector. It can be reliably distinguished from all other species of Serrivomer by its lower number of vertebrae (134 vs. 140–172) and its larger cleithrum, which is positioned under the seventh instead of the eighth vertebral centrum. http://zoobank.org/urn:lsid:zoobank.org:pub:50673E06-602C-4CA1-8CFF-AD992BCAEBED

An Eocene conger eel (Teleostei, Anguilliformes) from the Lillebælt Clay Formation, Denmark

A conger eel (Anguilliformes, Congridae) is described from the lower Lutetian concretionary nodules of the Lillebælt Clay Formation exposed at Trelde Næs, eastern Jutland, based on two partially complete articulated cranial skeletons. One of the cranial specimens exhibits an otolith void from which a cast was taken, used by Schwarzhans (2007) to describe the extinct Pseudoxenomystax treldeensis, which is placed herein within the new genus Smithconger gen. nov. Smithconger treldeensis (Schwarzhans, 2007) is characterized by well-developed lateral processes on the frontals, supraoccipital crest absent, sphenotic spine rather large, anteriorly pointed and exposed on the flattened surface of the skull roof, otic bullae considerably reduced, maxilla almost straight and distally pointed, maxillary and dentary teeth numerous and arranged in multiple rows, dentary with slightly convex ventral profile, opercle with smooth posterior margin and subopercle short. The otoliths of Smithconger treldeensis show high dorsal rim, broad and deep dorsal depression, no ventral furrow, sulcus straight, shallow, centrally positioned with anteriorly reduced colliculum, and ostial channel at anterior tip of colliculum short, not reaching the predorsal rim. The otolith-based species Bathycongrus waihaoensis Schwarzhans, 2019 from the Kaiatan (Bartonian/Priabonian) of New Zealand is also assigned to the genus Smithconger. Smithconger is tentatively referred to the congrid subfamily Congrinae due to the lack of hypohyals in the hyoid bar. This new Eocene genus of conger eel shows a certain degree of similarity with the extant Bassanago. The diversity and relationships of other Eocene congrids is also briefly discussed.

An Eocene conger eel (Teleostei, Anguilliformes) from the Lillebælt Clay Formation, Denmark

A conger eel (Anguilliformes, Congridae) is described from the lower Lutetian concretionary nodules of the Lillebælt Clay Formation exposed at Trelde Næs, eastern Jutland, based on two partially complete articulated cranial skeletons. One of the cranial specimens exhibits an otolith void from which a cast was taken, used by Schwarzhans (2007) to describe the extinct Pseudoxenomystax treldeensis, which is placed herein within the new genus Smithconger gen. nov. Smithconger treldeensis (Schwarzhans, 2007) is characterized by well-developed lateral processes on the frontals, supraoccipital crest absent, sphenotic spine rather large, anteriorly pointed and exposed on the flattened surface of the skull roof, otic bullae considerably reduced, maxilla almost straight and distally pointed, maxillary and dentary teeth numerous and arranged in multiple rows, dentary with slightly convex ventral profile, opercle with smooth posterior margin and subopercle short. The otoliths of Smithconger treldeensis show high dorsal rim, broad and deep dorsal depression, no ventral furrow, sulcus straight, shallow, centrally positioned with anteriorly reduced colliculum, and ostial channel at anterior tip of colliculum short, not reaching the predorsal rim. The otolith-based species Bathycongrus waihaoensis Schwarzhans, 2019 from the Kaiatan (Bartonian/Priabonian) of New Zealand is also assigned to the genus Smithconger. Smithconger is tentatively referred to the congrid subfamily Congrinae due to the lack of hypohyals in the hyoid bar. This new Eocene genus of conger eel shows a certain degree of similarity with the extant Bassanago. The diversity and relationships of other Eocene congrids is also briefly discussed.

Evolution and diversity of ram-suction feeding in damselfishes (Pomacentridae)

The cerato-mandibular (c-md) ligament is a synapomorphy within Pomacentridae that creates a tight link between the lower jaws and the hyoid bars. However, this morphological trait has been secondarily lost in multiple lineages during evolution. A previous study revealed that the loss of this trait acted as a release of evolutionary constraints, leading to a cascade of morphological changes such elongated buccal jaws and a slender body. Ecomorphological interpretations suggested the loss of the c-md ligament has ultimately led to a new adaptive peak in zooplanktivory through an optimization of the ram feeding mode associated with a specialization in pelagic feeding. Here, we tested these hypotheses by comparing functional and diet diversity between damselfish species with and without the c-md ligament. Although species lacking the c-md ligament presented a conserved kinematic pattern resulting from high ram and low suction performances, our results did not support an optimization of the ram feeding mode. Indeed, some species with the c-md ligament showed the same or exceeded the ram performance of species without the c-md ligament. The species with the c-md ligament had a more diverse kinematic pattern exploring the entire ram-suction functional range in damselfishes. Finally, our results did not show any diet variations associated with the loss of the c-md ligament. Our study furthers the understanding of how a morphological trait has shaped, by its presence or absence, the ecomorpho-functional diversification of Pomacentridae.

Insight into biting diversity to capture benthic prey in damselfishes (Pomacentridae)

The cerato-mandibular (c-md) ligament, joining the hyoid bar to the coronoid process of the angular, allows Pomacentridae to slam their mouth shut in a few milliseconds. Previous works have revealed that such a mechanism is used to feed, but some variability in biting patterns has been observed between two damselfish species. The pelagic feeder Amphiprion clarkii performs two different kinematic patterns to bite fixed prey, one that does not depend on the c-md ligament (biting-1) and one that does (biting-2). The benthic feeder Stegastes rectifraenum only performs biting-2. The present study aims to shed light on the occurrence of biting-2 in the feeding behaviour of Pomacentridae. To test our hypothesis that biting-2 would be the only biting pattern for benthic feeders, we compared biting behaviours among four species: one pelagic feeder, A. clarkii, and three benthic feeders, Neoglyphidodon nigroris, Stegastes leucostictus and S. rectifraenum. Our results showed that the four species were able to perform biting-2, but they do not support that the use of this pattern is related to trophic habits. Contrary to S. rectifraenum, the two other benthic feeders randomly used biting-1 and biting-2 patterns, similar to A. clarkii. Two hypotheses are discussed for explaining this variability within Pomacentridae. Finally, it has been recently shown that some damselfishes do not possess the c-md ligament. We therefore included two species lacking the c-md ligament (Chromis chromis and Abudefduf troschelii) in our study and we demonstrate our expectation that they are unable to perform biting-2.

The skull and endocranium of a Lower Jurassic ichthyosaur based on digital reconstructions

AbstractEven after 200 years of study, some details of the cranial anatomy of ichthyosaurs, one of the most successful groups of marine vertebrates in the Mesozoic, are still unclear. New information on the braincase, palate and occiput are provided from three‐dimensional scans of an exceptionally preserved ichthyosaur (‘Hauffiopteryx’ typicus) skull from the Toarcian (183–174 Ma, Lower Jurassic) of Strawberry Bank, England. This ichthyosaur has unusual, hollow, tubular hyoid bars. The occipital and braincase region is fully reconstructed, creating the first digital cranial endocast of an ichthyosaur. Enlarged optic lobes and an enlarged cerebellum suggest neuroanatomical adaptations that allowed it to be a highly mobile, visual predator. The olfactory region also appears to be enlarged, suggesting that olfaction was more important for ichthyosaurs than has been assumed. Phylogenetic analysis suggests this ichthyosaur is closely related to, but distinct from, Hauffiopteryx, and positioned within Thunnosauria, a more derived position than previously recovered. These results further our knowledge of ichthyosaur cranial anatomy in three dimensions and provide a platform in which to study the anatomical adaptations that allowed ichthyosaurs to dominate the marine realm during the Mesozoic.

A morphological novelty for feeding and sound production in the yellowtail clownfish.

The yellowtail clownfish Amphiprion clarkii is able to close its mouth very quickly by means of the cerato-mandibular (c-md) ligament, a synapomorphic trait of Pomacentridae joining the hyoid bar to the medial part of the lower jaw. This fast closure induces tooth collision, thus producing sounds that the clownfish uses during agonistic behaviors. To investigate whether this rapid jaw movement is also used during feeding, we analyzed the kinematics of sound production and feeding. Sound production, feeding on live planktonic prey, and feeding on food attached to tweezers was filmed with a high-speed camera. Three kinds of kinematic patterns were detected and were associated with the two different types of food presented: one performed to catch planktonic prey (PP), and two (called B-1 and B-2) to bite attached food items. The kinematic pattern of B-2 is similar to that observed during sound production (SP) and the transection of the c-md ligament highlights that sound production and biting-2 motions are dependent on this morphological trait. Our data show that the c-md ligament in addition to its role in sound production allows duplication of the mouth-closing mechanism during feeding. This highlights the key role played by the c-md ligament in sound production and feeding on attached prey. J. Exp. Zool. 323A: 227-238, 2015. © 2015 Wiley Periodicals, Inc.

The cerato-mandibular ligament: a key functional trait for grazing in damselfishes (Pomacentridae)

The success of a taxonomic group can be promoted by a key character that allows the group to interact with its environment in a different way and to potentially occupy new niches. The Pomacentridae possess a synapomorphic trait, the cerato-mandibular (c-md) ligament, which joins the hyoid bar to the inner part of the lower jaw. It has previously been shown that this ligament is a key trait in communication in damselfishes because it enables them to slam the oral jaws shut causing teeth collision and sound production. This specific behavior of mouth closing could, however, also be used for other tasks, such as feeding. Many territorial damselfishes are referred to as farmers, due to their ability to manage algal crops on which they feed. This study hypothesizes that the c-md ligament provides an advantage for grazing filamentous algae, and should thus be considered a key trait for farming behavior. The kinematic patterns associated with sound production and biting filamentous algae or attached animal prey are all based on the same mechanism and are associated with a slam of the oral jaws. We observed that transection of the c-md ligaments makes the fish unable to perform such actions. We also counted biting rates on filamentous algae in fish with and without the c-md ligament and observed a drop of more than 80% in the latter. This study shows that the c-md ligament is a key trait both for sound production and for grazing activities in damselfishes. The buccal jaw slam enables the fish to perform accurate strikes on small filamentous algae. This kind of bite probably plays a major role in farming activity and allows grazing damselfishes to occupy distinct niches, possibly increasing their competitive evolutionary success.

Trait decoupling promotes evolutionary diversification of the trophic and acoustic system of damselfishes.

Trait decoupling, wherein evolutionary release of constraints permits specialization of formerly integrated structures, represents a major conceptual framework for interpreting patterns of organismal diversity. However, few empirical tests of this hypothesis exist. A central prediction, that the tempo of morphological evolution and ecological diversification should increase following decoupling events, remains inadequately tested. In damselfishes (Pomacentridae), a ceratomandibular ligament links the hyoid bar and lower jaws, coupling two main morphofunctional units directly involved in both feeding and sound production. Here, we test the decoupling hypothesis by examining the evolutionary consequences of the loss of the ceratomandibular ligament in multiple damselfish lineages. As predicted, we find that rates of morphological evolution of trophic structures increased following the loss of the ligament. However, this increase in evolutionary rate is not associated with an increase in trophic breadth, but rather with morphofunctional specialization for the capture of zooplanktonic prey. Lineages lacking the ceratomandibular ligament also shows different acoustic signals (i.e. higher variation of pulse periods) from others, resulting in an increase of the acoustic diversity across the family. Our results support the idea that trait decoupling can increase morphological and behavioural diversity through increased specialization rather than the generation of novel ecotypes.

A novel pharyngeal expansion mechanism in the yellow-spotted fanray, Platyrhina tangi (Elasmobranchii: Batoidea), with special reference to the function of the fifth ceratobranchial cartilage in batoids

Expansion of the ‘pharynx’ during breathing or capturing prey in fishes generally involves posteroventral retraction of the hyoid arch. However, the hyoid arch structure of batoid fishes (skates, rays, guitarfishes, and sawfishes) is unique, and how they expand the pharyngeal cavity is poorly understood. To investigate the mechanism of pharyngeal expansion during breathing in the yellow-spotted fanray, Platyrhina tangi, we conducted anatomical and kinematic investigations of the pharyngeal region. Our study revealed that the yellow-spotted fanray and sharks have different skeletal linkage systems for pharyngeal expansion. During pharyngeal expansion in the yellow-spotted fanray, the hyoid bar and branchial apparatus rotate ventrally around the hinge joint between the fifth ceratobranchial cartilage and the pectoral girdle. This pharyngeal expansion mechanism appears to be widespread among batoid fishes and is unique among cartilaginous fishes (sharks, batoids, and holocephalans). Batoid fishes possibly developed this pharyngeal expansion mechanism during early batoid evolution.

Mechanics of snout expansion in suction feeding seahorses: musculoskeletal force transmission

Seahorses and other syngnathid fishes rely on a widening of the snout to create the buccal volume increase needed to suck prey into the mouth. This snout widening is caused by abduction of the suspensoria, the long and flat bones outlining the lateral sides of the mouth cavity. However, it remains unknown how seahorses can generate a forceful abduction of the suspensoria. To understand how force is transmitted to the suspensoria via the hyoid and the lower jaw, we performed mathematical simulations with models based on computerized tomography scans of Hippocampus reidi. Our results show that the hinge joint between the left and right hyoid bars, as observed in H. reidi, allows for an efficient force transmission to the suspensorium from a wide range of hyoid angles, including the extremely retracted hyoid orientations observed in vivo for syngnathids. Apart from the hyoid retraction force by the sternohyoideus-hypaxial muscles, force generated in the opposite direction on the hyoid by the mandibulohyoid ligament also has an important contribution to suspensorium abduction torque. Forces on the lower jaw contribute only approximately 10% of the total suspensorium torque. In particular, when dynamical aspects of hyoid retraction are included in the model, a steep increase is shown in suspensorium abduction torque at highly retracted hyoid positions, when the linkages to the lower jaw counteract further hyoid rotation in the sagittal plane. A delayed strain in these linkages allows syngnathids to postpone suction generation until the end of cranial rotation, a fundamental difference from non-syngnathiform fishes.

Biting releases constraints on moray eel feeding kinematics

We present an analysis of prey capture functional morphology in eels by comparing two species of moray eels, Muraena retifera and Echidna nebulosa (Family Muraenidae), to the American eel Anguilla rostrata (Family Anguillidae). The skulls of both moray species exhibited extreme reductions of several prominent components of the suction-feeding mechanism, including the hyoid bar, the sternohyoideus muscle and the pectoral girdle. Associated with these anatomical modifications, morays showed no evidence of using suction during prey capture. From 59 video sequences of morays feeding on pieces of cut squid we saw no hyoid depression and no movement of prey toward the mouth aperture during the strike, a widely used indicator of suction-induced water flow. This was in contrast to A. rostrata, which exhibited a robust hyoid, sternohyoideus muscle and pectoral girdle, and used suction to draw prey into its mouth. Average prey capture time in morays, about 500 ms, was roughly 10 times longer than in A. rostrata, and morays frequently reversed the direction of jaw and head rotation in the midst of the strike. We tested whether the absence of suction feeding reduces temporal constraints on feeding kinematics, permitting greater variance in traits that characterize timing and the extent of motion in the neurocranium, by comparing moray eel species with A. rostrata, two Centrarchids and a cichlid. Kinematic variance was roughly 5 times higher in morays than the suction-feeding species. Prey capture by suction demands a rapid, highly coordinated series of cranial movements and the loss of this mechanism appears to have permitted slower, more variable prey capture kinematics in morays. The alternative prey capture strategy in morays, biting, may be tied to their success as predators in the confined spaces of reef crevices where they hunt for cephalopods, crustaceans and fish.

Early development of the chondrocranium in Chrysichthys auratus

The inception and development of the cartilaginous cephalis skeleton of Chrysichthys auratus is described from hatching to about 18 days post‐hatching. At hatching, no skeletal structure is present. Not until day 3 do clearly delimited cranial primordia become apparent. As in many siluriforms, the neurocranium is platybasic from the start, the suspensorium constitutes, with Meckel's cartilage and the hyoid bar, a single cartilaginous element, and the junction between the front and rear of the neurocranium is complete on day 4. By day 8 the quadratomandibular joint has formed and the tectum posterius has appeared. Cartilage reduction first affects the trabecular bars, then, markedly, the visceral arches. By day 18 the braincase floor has almost disappeared.

Development of the cartilaginous skull in solea solea: trends in pleuronectiforms

The postembryonic development of the cartilaginous cephalic skeleton in Solea solea (Linné) was observed from hatching to the juvenile stage or postmetamorphic larva. Fry were trypsin-cleared and alcian-blue-stained. At hatching, Solea solea displays no cartilaginous structure. On day 3, the fry exhibit trabecular bars, Meckel's cartilages, palatoquadrates, hyosymplectics, hyoid bars, the first two ceratobranchials, and basibranchials 1–3. On day 4, ceratobranchials 3 and 4 form. On day 12, short taeniae marginales are formed on the well-developed otic capsules and the branchial basket is nearly complete. On day 14, a first cartilage reduction begins as the posterior part of the trabecula communis separates from the otic capsules. On day 18, metamorphosis begins. The tectum posterius completes posteriorly the braincase dome. On day 20, the left eye moves to the median crest of the head and the right eye shifts downwards slightly. Consequently, displacement of the right eye induces a deformation extending to the right pterygoid process. On day 23, both eyes are located on the same side. On day 29, cartilage reduction affects the hyosymplectics; it continues with the first two ceratobranchials on day 32 and with the otic capsules on day 40. On day 50, cartilage reduction has reached the lamina precerebralis. Only a few small lumps remain at the level of the neurocranium and the splanchnocranium.

Ontogeny of the osteocranium in the African catfish, Clarias gariepinus Burchell (1822) (Siluriformes: Clariidae): Ossification sequence as a response to functional demands.

The ontogeny of the bony skull of the African catfish, Clarias gariepinus, is studied from initial ossification until a complete skull is formed. The ossification sequence in C. gariepinus seems to be related to the functional demands that arise in a developing larva. Early ossification of the opercular bone coincides with the initiation of opercular skin movements. Early ossifications involve several dentulous bones, formed shortly before the transition phase from endogenous to exogenous feeding. The enlarging branchiostegal membrane becomes supported by the gradual adding of branchiostegal rays. Parasphenoid ossification may be related to protection of the brain during prey transport, whereas the several hyoid bones, including the parurohyal, are formed in relation to the increasing loads exerted onto the tendons of the sternohyoideus and consequently onto the hyoid bar. Overall skull reinforcement occurs almost simultaneously, with a whole set of perichondral bones arising especially at places of high mechanical load. The suspensorium becomes protected against dislocation in an anteroposterior direction through a ligamentous connection, which even becomes partially ossified, forming the sesamoid entopterygoid. Later, the cranial lateral-line system becomes enclosed by a set of gutters, which close, frequently becoming plate-like later in ontogeny. The brain also becomes covered dorsally. Additional dentition (prevomeral tooth plates) formation seems to coincide with formation of the opercular four-bar system, as well as with the time the digestive system becomes completely functional. Eventually, unossified regions between the bones become closed off, fortifying and completely covering the skull. J. Morphol. 235:183-237, 1998. © 1998 Wiley-Liss, Inc.

Early development of the cephalic skeleton in the turbot

At hatching Scophthalmus maximus shows no cartilaginous and no bony structure. Meckeľs cartilages appear when the fry are 1 day old, followed on day 2, by formation of the trabecular bars, fused at the outset to form a trabecula communis. Concurrently, the palatoquadrates complete the mandibular arch, and the first two pairs of ceratobranchials, associated with a pair of hyoid bars, form the beginnings of the hyobranchial system. By day 3, the parachordals have fused with the trabecular bars, the hyosymplectics have linked to the hyoid bars by interhyals, and the first four pairs of ceratobranchials have appeared. The first bony structures appear: the preoperculars. On day 8, the frontals develop above the orbits and the maxillaries and dentaries appear. On day 10, the primordia of the taeniae marginales appear, the palatoquadrates bear a pterygoid process, and to the branchial basket have been added the fifth pair of ceratobranchials and the four pairs of epibranchials. On day 12, both pairs of posterior pharyngobranchials are present. The premaxillaries develop in front of the maxillaires, and retroarticulars and the angulars complete the lower jaws. On day 13, a thin parasphenoid contributes to the floor of the neurocranium, and ectopterygoids and entopterygoids to the splanchnocranium. The set of opercular bones is complete. On day 15, the tectum synoticum closes the braincase posteriorly. The splanchnocranium possesses a basihyal and the pharyngobranchials of the first epibranchials. On day 18, the tectum posterius completes the dome of the braincase. The rear end and lateral walls of the skull are formed by the basioccipital, the exoccipitals, the pterotics, and the parietals. The suspensorium is nearly complete. From day 10, the first resorptions begin in parallel with the construction of the chondrocranium. Meckeľs cartilages each split in two, then the posterior part of the trabecular bars disappears. On day 23, the right taenia marginalis separates from the lamina orbitonasalis and curves towards the centre. Simultaneously, the right eye begins its migration to the left. This is the only metamorphosis-linked asymmetry to appear during the development of the chondrocranium. On day 25, many more bony structures appear, a characteristic of this stage: the nasals, lateral ethmoids, mesethmoid, sphenotics, prootics, pleurosphenoids, epiotics, and supraoccipital. From this stage on, the bony structures continue to develop, while the front of the neurocranium and the jaws undergo a deep remodelling due to metamorphosis. The left taenia marginalis does not appear reduced until day 29. By day 45, there remain only a few small elements of the cartilaginous skull.

Early development of the cephalic skeleton in the turbot

At hatching Scophthalmus maximus shows no cartilaginous and no bony structure. Meckeľs cartilages appear when the fry are 1 day old, followed on day 2, by formation of the trabecular bars, fused at the outset to form a trabecula communis. Concurrently, the palatoquadrates complete the mandibular arch, and the first two pairs of ceratobranchials, associated with a pair of hyoid bars, form the beginnings of the hyobranchial system. By day 3, the parachordals have fused with the trabecular bars, the hyosymplectics have linked to the hyoid bars by interhyals, and the first four pairs of ceratobranchials have appeared. The first bony structures appear: the preoperculars. On day 8, the frontals develop above the orbits and the maxillaries and dentaries appear. On day 10, the primordia of the taeniae marginales appear, the palatoquadrates bear a pterygoid process, and to the branchial basket have been added the fifth pair of ceratobranchials and the four pairs of epibranchials. On day 12, both pairs of posterior pharyngobranchials are present. The premaxillaries develop in front of the maxillaires, and retroarticulars and the angulars complete the lower jaws. On day 13, a thin parasphenoid contributes to the floor of the neurocranium, and ectopterygoids and entopterygoids to the splanchnocranium. The set of opercular bones is complete. On day 15, the tectum synoticum closes the braincase posteriorly. The splanchnocranium possesses a basihyal and the pharyngobranchials of the first epibranchials. On day 18, the tectum posterius completes the dome of the braincase. The rear end and lateral walls of the skull are formed by the basioccipital, the exoccipitals, the pterotics, and the parietals. The suspensorium is nearly complete. From day 10, the first resorptions begin in parallel with the construction of the chondrocranium. Meckeľs cartilages each split in two, then the posterior part of the trabecular bars disappears. On day 23, the right taenia marginalis separates from the lamina orbitonasalis and curves towards the centre. Simultaneously, the right eye begins its migration to the left. This is the only metamorphosis‐linked asymmetry to appear during the development of the chondrocranium. On day 25, many more bony structures appear, a characteristic of this stage: the nasals, lateral ethmoids, mesethmoid, sphenotics, prootics, pleurosphenoids, epiotics, and supraoccipital. From this stage on, the bony structures continue to develop, while the front of the neurocranium and the jaws undergo a deep remodelling due to metamorphosis. The left taenia marginalis does not appear reduced until day 29. By day 45, there remain only a few small elements of the cartilaginous skull.

Ontogeny of the hyoid musculature in the African catfish,Clarias gariepinus(Burchell, 1822) (Siluroidei: Clariidae)

Three ontogenetic stages of the African catfishClarias gariepinushave been used to describe and discuss the ontogeny of the hyoid musculature. During ontogeny, an asynchrony in the development of the muscles is observed: the intermandibularis and protractor hyoidei are the first to develop and which bear their insertions, followed by the hyohyoideus inferior and the sternohyoideus. The hyohyoideus abductor and adductor muscles are the last of the hyoid muscles to develop. In the juvenile stage (136.2 mm SL specimen), the intermandibularis is still present. The protractor hyoidei is well developed, as it may play an important role in the opening of the mouth, the elevation of the hyoid bars and, as a typical catfish feature, the displacement of the mandibular barbels. The protractor hyoidei arises as three pairs of muscle bundles (a pars ventralis, a pars lateralis and a pars dorsalis), of which the pars ventralis and the pars lateralis become fused to each other. This fusion gives rise to four different fields of superficial fibres for the manipulation of the mandibular barbels. The pars dorsalis, with its tendinous insertion, may be of more importance for mouth opening and/or hyoid elevation. The hyohyoid muscle is well differentiated into an inferior, abductor and adductor muscles, acting on the hyoid bars, the branchiostegal rays and the opercular bone.

Ontogeny of the hyoid musculature in the African catfish, Clarias gariepinus (Burchell, 1822) (Siluroidei: Clariidae)

Abstract Three ontogenetic stages of the African catfishClarias gariepinushave been used to describe and discuss the ontogeny of the hyoid musculature. During ontogeny, an asynchrony in the development of the muscles is observed: the intermandibularis and protractor hyoidei are the first to develop and which bear their insertions, followed by the hyohyoideus inferior and the sternohyoideus. The hyohyoideus abductor and adductor muscles are the last of the hyoid muscles to develop. In the juvenile stage (136.2 mm SL specimen), the intermandibularis is still present. The protractor hyoidei is well developed, as it may play an important role in the opening of the mouth, the elevation of the hyoid bars and, as a typical catfish feature, the displacement of the mandibular barbels. The protractor hyoidei arises as three pairs of muscle bundles (a pars ventralis, a pars lateralis and a pars dorsalis), of which the pars ventralis and the pars lateralis become fused to each other. This fusion gives rise to four different fields of superficial fibres for the manipulation of the mandibular barbels. The pars dorsalis, with its tendinous insertion, may be of more importance for mouth opening and/or hyoid elevation. The hyohyoid muscle is well differentiated into an inferior, abductor and adductor muscles, acting on the hyoid bars, the branchiostegal rays and the opercular bone.

Postembryonic development of the cephalic region in Heterobranchus longifilis

At hatching, Heterobranchus longifilis does not display any primordia of the cephalic skeleton. The latter appears 12 h post–hatching and develops in three stages up to day 16. The first stage (12 h to 2 days) involves almost exclusively the development of the chondrocranium. During the second period (days 3–8), dermal elements of the splanchnocranium appear. The final stage is marked by resorption of the cartilages, progressively replaced by ossifications (days 10–16). At their appearance the elements of the splanchnocranium are fused together, as are the first neurocranial elements. Later, the splanchnocranium splits up. By the time the yolk sac is completely resorbed, the buccal and pharyngeal jaws are present, the suspensoria and hyoid bars are partially developed, and the parasphenoid partially closes the hypophyseal fenestra. These structures delimit a buccal cavity that is probably functional, i.e. capable of participating in the intake of exogenous food. Next to continue its development is principally the splanchnocranium, completing the walls of the buccal cavity. Cartilage resorption parallels the appearance of endochondral ossifications (except for the trabecular bars). Braincase closure begins to accelerate once the buccal system is complete.