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.     

The ontogeny of the chondrocranium in Clarias gariepinus: trends in siluroids

The ontogeny of the chondrocranium of 31 different stages of the African catfish Clarias gariepinus (Siluroidei: Clariidae) was studied, both from cleared and stained, and sectioned material. The fish ranged from 4.1 (1 day post-hatching) to 127.0 mm SL (100 days post-hatching). The chondrocranium of C. gariepinus seemed to correspond to the general adaptive trends in siluroids, especially in relation to the reduction of eye size and the dorso-ventral flattening of the skull. The platybasic neurocranium involved several modifications related to the trabecular bars, the hypophyseal fenestra, the ethmoid region and even the olfactory nerves. Certain reductions were present, which have been observed in all siluroids (e.g. absence of the pila lateralis, the commissura lateralis, the myodomes) or are part of a variable trend within siluroids (e.g. reduction of the taenia marginalis anterior and the tectum synoticum). Compared with some other siluroid species, the neurocranium of C. gariepinus is well developed, for example in the otic region. The same was observed in the splanchnocranium where some general siluroid trends persist (e.g. isolation of palatine from pterygoquadrate, presence of 'hyo-symplectic-pterygoquadrate' plate). Some trends, as observed in other siluroids, were present also (e.g. interhyal continuous with suspensorium and ceratohyal, Meckel's cartilage initially continuous with the suspensorium). The branchial basket is well developed as all expected elements are present (basibranchials I-IV, hypobranchials I-IV, ceratobranchials I-V, epibranchials I-IV). Based on the observed ontogeny of C. gariepinus and data from the literature, a hypothesis was formulated which indicated the presence of a general reductional trend within siluroids. In C. gariepinus , all four (I-IV) infrapharyngobranchials develop, although the anterior two are much reduced and fused with each other.     

Early development of the cephalic skeleton of Barbus barbus (Teleostei, Cyprinidae)

The inception, and development of the cephalic skeleton of Barbus barbus from hatching to 24 days passes through periods of fast and slow growth; these rates are not the same in different parts of the skull. Trabeculae, parachordal plates, Meckelian cartilages and hyposymplectics are present at hatching. Then the cartilaginous floor of the neurocranium develops, the pars quadrata, the hyoid bars and branchial arches elements appear shortly before the first movable dermal bones, the dentaries, maxillae and opercles. The first bone of the braincase to appear is the parasphenoid; other bones develop subsequently and at the same time: the angular, quadrate, interopercle and fifth ceratobranchial. Later the splanchnocranium continues to develop at a relatively fast rate while the neurocranium shows little growth. The braincase does not begin to close before the 24th day, nor do the first bones of the skull roof appear, while the bucco-pharyngeal apparatus is complete, having the adult shape. The early constitution of the latter structures seems to be linked with the mechanical demands of biological functions such as breathing and feeding.     

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.     

Studies on the morphology of the gills and gill arches with reference to the blood supply in Notopterus motopterus and Colisa fasciatus.

The morphology of the gills, with their blood supply have been described in Notopterus notopterus and Colisa fasciatus in some detail. Gills are curved and perforated on the dorsolateral and ventrolateral wall of the pharynx. The gills consist of 2 rows of filaments which are stacked one above the other to form a space. The gill filaments are smaller on both the ends and larger in middle. The gill filaments are of pink colour as they are supplied with blood. Gill rakers are large in size in Notopterus notopterus while they are small in Colisa fasciatus. 3 pairs of basibranchials are present in Notopterus notopterus which are covered by median membranous bony plate while 2 basibranchials are present in Colisa fasciatus. 3 pairs of hypobranchials are present in both fishes. 5 pairs of ceratobranchials are present in which Vth ceratobranchial bears teeth. 4 pairs of epibranchials are present. 3 pairs of pharyngobranchials are present in which the tip of the IVth pharyngobranchial bears minute teeth in Notopterus notopterus while in Colisa fasciatus IInd and IIIrd pharyngobranchial bear minute ones. One afferent branchial vessel is present in Notopterus notopterus and Colisa fasciatus in each gill like in other teleostean fishes. One efferent branchial vessel is present in each gill of Notopterus notopterus while in Colisa fasciatus 2 efferent are represented in each gill.     

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.     

Atavisms and the homology of hyobranchial elements in lower vertebrates

The homology of branchial arch segments in salamanders has been a matter of controversy since the last century. Many investigators term the most medial paired elements of salamander branchial arches “ceratobranchials” and the next distal paired elements “epibranchials.” This suggests that the first two segmental elements of the salamander branchial arch are not homologous with elements occupying the same position in ray-finned fishes, Latimeria, “rhipidistians,” and lungfishes, in which these bones are called hypobranchials and ceratobranchials, respectively. Three lines of evidence suggest that it is more parsimonious to interpret urodele branchial arch segments as being homologous with those of other vertebrate clades?(1) comparative osteology, (2) comparative myology, and (3) the discovery of cartilaginous structures forming a third segmental unit that we interpret as atavistic epibranchials of the branchial arch in one population of the salamander Notophthalmus viridescens. These structures possess all the defining attributes of atavisms, and illustrate the special role that atavistic features play in resolving questions of homology recognition.     

Cranial skeletogenesis and osteology of the redeye tetra Moenkhausia sanctaefilomenae

The skeletogenesis and osteology of the syncranium of the redeye tetra Moenkhausia sanctaefilomenae is described. Skeletal development is rapid, with many elements of the chondrocranium and splanchnocranium well formed prior to the onset of ossification. The chondrocranium develops from an initial set of cartilaginous precursors, and continued elaboration proceeds from a series of processes which expand and converge to form the floor of the cranial vault, the otic capsule, the supraorbital bridge and the ethmoid region. Prodigious growth is observed for a number of splanchnocranial elements, including the Meckel's cartilage and the ceratohyal cartilage. Ossification occurs in overlapping phases with initial ossification of the jaws and neurocranial floor followed by the splanchnocranium, the supraorbital bridges and the ethmoid and cranial vault. Teeth are observed primarily on the premaxilla and dentary, while a single tooth is present on the maxilla. Particular cartilages, which had originally formed in the early larva, appear to degenerate and have no ossified representative in the adult syncranium. The cranial development for M. sanctaefilomenae is compared to those of other characiforms.     

A complex hyobranchial apparatus in a Cretaceous dinosaur and the antiquity of avian paraglossalia

The highly specialized feeding apparatus of modern birds is characterized in part by paraglossalia, triangular bones or cartilages in the tongue that constitute part of the rarely fossilized hyobranchial apparatus. Here, we report on a new, juvenile specimen of the ankylosaurid dinosaur Pinacosaurus grangeri Gilmore, 1933 that provides the first evidence of paraglossalia outside of crown group Aves. The specimen is remarkable in preserving a well‐ossified hyobranchial apparatus, including paired paraglossalia, first and second ceratobranchials, epibranchials, and evidence of a median cartilaginous basihyal. Reassessment of Edmontonia, another ankylosaur, also reveals evidence of bony paraglossalia. Ankylosaur paraglossalia closely resemble those of birds, but are relatively larger and bear prominent muscle scars, supporting the hypothesis that ankylosaurs had fleshy, muscular tongues. The other hyobranchial elements, surprisingly, resemble those of terrestrially feeding salamanders. Ankylosaurs had reduced, slowly replacing teeth, as evidenced from dental histology, suggesting that they relied greatly on their tongues and hyobranchia for feeding. Some curved, rod‐like elements of other dinosaur hyobranchia are reinterpreted as second ceratobranchials, rather than first ceratobranchials as commonly construed. Ankylosaurs provide rare fossil evidence of deep homology in vertebrate branchial arches and expose severe biases against the preservation and collection of the hyobranchial apparatus. In light of these biases, we hypothesize that paraglossalia were present in the common ancestor of Dinosauria, indicating that some structures of the highly derived avian feeding apparatus were in place by the Triassic Period. © 2015 The Linnean Society of London     

Development of the splanchnocranium in Prochilodus argenteus (Teleostei: Characiformes) with a discussion of the basal developmental patterns in the Otophysi

Development of the mandibular, hyoid and gill arches, which constitute the splanchnocranium, are described for Prochilodus argenteus, order Characiformes, one of the basal lineages of the Otophysi. Development was examined from just hatched larvae through juveniles using whole specimens cleared and counterstained for cartilage and bone as well as histological preparations. Observations are compared with the developmental trends reported for Cypriniformes, the basalmost clade of the Otophysi. Shortened developmental sequences for Prochilodus compared to the cypriniform Catostomus were discovered in the ontogeny of the ceratohyals, ceratobranchials 1–5, epibranchials 1–4 and the symplectic portion of the hyosymplectic. Prochilodus also differs from Catostomus in having the basihyal plus the anterior copula appearing at different stages of ontogeny rather than simultaneously. Contrary to previous assumptions, developmental information indicates that hypobranchial 4 as well as likely basibranchial 5 are present in Prochilodus. Various developmental patterns in Prochilodus considered basal for the Otophysi, the predominant component of the Ostariophysi, are likely conserved from patterns prevalent in basal groups in the Actinopterygii.     

Early development of the chondrocranium in Salmo letnica(Karaman, 1924)(Teleostei: Salmonidae)

The ontogenetic development of the chondrocranium of Ohrid trout Salmo letnica was studied from hatching until 92 days post‐hatching (dph). Most of the samples were in toto trypsin cleared and stained, some specimens were used for serial histological sectioning. The serial histological sections of fish specimens at the age of 92 dph were used for a graphical reconstruction of the cartilaginous neurocranium. A chronological evaluation of the formation of the cartilaginous skull in the early development of S. letnica was performed. In order to investigate to what degree the ontogeny of the Ohrid trout is unique, the results were compared with data of the development of other salmonids, as well as some non‐salmonid teleosts. The development of the cartilaginous structures of the Ohrid trout was found to be similar to that of other salmonids. Most of the cartilage structures of the neurocranium and the viscerocranium are present at the moment of hatching of this species. A fully developed chondrocranium was observed at the age of 92 dph, when the first signs of cartilage resorption could also be observed.     

Development of the bony skull in common sole: brief survey of morpho-functional aspects of ossification sequence

The postembryonic development of the bony cephalic skeleton in the common sole Solea solea , observed from hatching to the juvenile stage or postmetamorphic larva, appears to follow a similar chronological order to that observed in other Pleuronectiformes and Perciformes and the sequence in bone formation is a response to functional demands. At hatching, S. solea has no bony structure. On day 4, only the outlines of maxillaries and opercular bones are visible. On day 6, a thin parasphenoid appears between the orbits and isolates the braincase from the buccal cavity making food ingestion possible without any impact on the brain. On day 8, the dentaries form and two small preopercular bones appear on each side of the head. On day 9, at weaning from the yolk sac, branchial arches support the gill filaments (used for respiration and trapping phytoplankton which pass through the open mouth). On day 10, the premaxillaries develop in front of the maxillaries. The superimposing of the maxillaries and the premaxillaries is a typical feature of species possessing an acanthopterygian protractile mouth at the adult stage. On day 12, the frontals develop above the orbits and the set of opercular bones is complete. On day 18, the migration of the left eye begins. On day 20, the left eye has moved to the median crest of the head. On day 23, both eyes are located on the same side. On day 26, the braincase is formed by a basioccipital, exoccipitals, pterotics, sphenotics and a supraoccipital. On day 50, new structures have appeared, others have developed and have undergone an extensive remodeling due to metamorphosis.     

鲂鱼的头骨发育及其适应意义

本文对鲂鱼（Ｍｅｇａｌｏｂｒａｍａｓｋｏｌｋｏｖｉｉ）头骨的早期发育过程及其与鱼苗的存活功能需要之间的关系进行了研究。头骨发育的全过程可划分为５个阶段,即软颅阶段、咽颅膜骨附加阶段、脑颅开始骨化阶段、脑颅快速骨化阶段和骨化完成阶段。刚出膜仔鱼头部即有软骨存在,最先出现的硬骨是膜质上颌骨和主鳃盖骨,脑颅最先开始骨化的是基枕骨和侧枕骨,随后才是副蝶骨。头骨发育过程与鱼苗早期存活的功能需要之间有着密切的关系。     

Covariation of brain and skull shapes as a model to understand the role of crosstalk in development and evolution

Covariation among discrete phenotypes can arise due to selection for shared functions, and/or shared genetic and developmental underpinnings. The consequences of such phenotypic integration are far-reaching and can act to either facilitate or limit morphological variation. The vertebrate brain is known to act as an “organizer” of craniofacial development, secreting morphogens that can affect the shape of the growing neurocranium, consistent with roles for pleiotropy in brain–neurocranium covariation. Here, we test this hypothesis in cichlid fishes by first examining the degree of shape integration between the brain and the neurocranium using three-dimensional geometric morphometrics in an F₅ hybrid population, and then genetically mapping trait covariation using quantitative trait loci (QTL) analysis. We observe shape associations between the brain and the neurocranium, a pattern that holds even when we assess associations between the brain and constituent parts of the neurocranium: the rostrum and braincase. We also recover robust genetic signals for both hard- and soft-tissue traits and identify a genomic region where QTL for the brain and braincase overlap, implicating a role for pleiotropy in patterning trait covariation. Fine mapping of the overlapping genomic region identifies a candidate gene, notch1a, which is known to be involved in patterning skeletal and neural tissues during development. Taken together, these data offer a genetic hypothesis for brain–neurocranium covariation, as well as a potential mechanism by which behavioral shifts may simultaneously drive rapid change in neuroanatomy and craniofacial morphology.     

Early development of the head skeleton in Brycon moorei(Pisces, Ostariophysi, Characidae)

At hatching (15 h post fertilization), Brycon moorei possesses no skeletal structure. Thereafter, development is very rapid. The first oral teeth appear no later than 3 h post‐hatching, but they remain covered with epithelium until c . 45 h. At 7 h, the trabecular bars and part of the cartilaginous visceral arches are visible and at 15 h, the dentaries and premaxillaries are present. At 25 h, i.e . the onset of piscivory and cannibalism (the yolk sac is only fully resorbed after 36 h), the oral teeth are fully developed, the first pharyngeal teeth are formed, and some head movements already appear synchronized, but the mouth cavity is not completely isolated from the neurocranium by bony structures. Thereafter, no new buccal or pharyngeal bony structure is visible until 45 h, when the maxilla and opercula appear, along with a new type of cannibalistic behaviour. Cartilage resorptions also start at 45 h, but with no concomitant replacement by formation of calcified structures. Later, development gradually becomes similar to that of many previously studied teleosts. The developmental pattern of B. moorei is thus extremely rapid in comparison with other teleosts, i.e . it prioritizes feeding structures that permit the expression of piscivory at a very early age. The uniqueness of this pattern is discussed in relation to ecological constraints on early feeding and fast growth.     

The braincase of the Middle Triassic shark Acronemus tuberculatus (Bassani, 1886)

Abstract: The chondrocranium of the enigmatic Middle Triassic shark Acronemus tuberculatus is investigated using computerized tomography scanning and 3‐D digital reconstitution techniques. The braincase reveals some autapomorphies, plus other features that suggest a phylogenetic relationship to both hybodontiform and neoselachian elasmobranchs, including evidence of features implicated in low‐frequency semi‐directional phonoreception. Acronemus can no longer be classified as a ‘ctenacanthiform’, although its relationships remain elusive and it presents supposedly hybodontiform and neoselachian features that have not previously been found in combination.     

Major stages in the evolution of primate neurocrania

An important constraint on the evolution of primate skeletons is the isometric relationship which exists between skeletal weight and body weight. The evolution of primate skulls during the Tertiary and Quaternary periods indicates that redeployment of bone mass took place largely within the skull (i.e. between proximate ossifying centres) and that the major vector was from the splanchnocranium towards the neurocranium. This vector of bone mass redeployment accords well with the general treand within primates of increased encephalisation over time. There are however, several interesting examples of vectors which were oriented in the opposite sense, in particular in the robust australopithecine lineage, in which the emphasis on bone mass deployment was towards the splanchnocranium and away from the neurocranium. A fuller understanding of skeletal isometry, and a wider application in comparative anatomy may throw much light on the evolution of skeletal systems, and it may resolve somelong-standing debates. Among these may be identification of the selection pressures which have led to dental and alveolar reduction inHomo sapiens sapiens (bone mass redeployment into the neurocranium) and perhaps an explanatation for some types of osteoporosis in old people whose body weight decreases may result in isometric skeletal mass decreases (for every 100 gms muscle tissue loss, there will be about 7 gms of bone tissue loss).     

Morphometric variation of the skull during postnatal development in the Lowland European bisonBison bonasus bonasus

We studied the variation of linear measurements and skull capacity in Lowland European bisonBison bonasus bonasus (Linnaeus, 1758) during postnatal development, and the dependencies of the parameters in relation to sex, age, and body mass of the animals. Material consisted of 599 bison skulls (310 males and 289 females), within the age range of 1 month to 21 years (males) and to 27 years (females). In the group of calves to 1 year old, no sex connected differences in skull measurements were observed, whereas the skull capacity in older calves was significantly larger (0.01>p>0.001) in males than in females. From the third year of life, most skull measurements display characteristics of sexual dimorphism. Skull development in both sexes is most intensive during the first three years of life, and slows from the age of 5. In older individuals of both sexes (≥ 6 years), orbital breadth continues growing and, in females, breadth of splanchnocranium continues increasing. Growth in a bison’s skull capacity is most intensive up to the third year of life and slows from the age of 5. During postnatal development, a bison skull grows proportionally except the neurocranium, which grows slightly slower in comparison with basal length and its development finishes earlier than that of splanchnocranium. In ontogenesis, a bison skull grows much slower compared to body mass. In relation to body mass, skull capacity and the height of neurocranium grow most slowly while orbital breadth grows most intensively. The results obtained were compared with data on skull sizes of bison born in 1930–1950 and bred in captivity and with skulls of the American bisonBison bison. Inbreeding is probably responsible for some types of phenotypic abnormalities in the skull which appear in modern European bison.