Histochemical study of jaw muscle fibers in the American alligator (Alligator mississippiensis)

The ontogenetic development of caudal vertebrae and associated skeletal elements of salmonids provides information about sequence of ossification and origin of bones that can be considered as a model for other teleosts. The ossification of elements forming the caudal skeleton follows the same sequence, independent of size and age at first appearance. Dermal bones like principal caudal rays ossify earlier than chondral bones; among dermal bones, the middle principal caudal rays ossify before the ventral and dorsal ones. Among chondral bones, the ventral hypural 1 and parhypural ossify first, followed by hypural 2 and by the ventral spine of preural centrum 2. The ossification of the dorsal chondral elements starts later than that of ventral ones. Three elements participate in the formation of a caudal vertebra: paired basidorsal and basiventral arcocentra, chordacentrum, and autocentrum; appearance of cartilaginous arcocentra precedes that of the mineralized basiventral chordacentrum, and that of the perichordal ossification of the autocentrum. Each ural centrum is mainly formed by arcocentral and chordacentrum. The autocentrum is irregularly present or absent. Some ural centra are formed only by a chordacentrum. This pattern of vertebral formation characterizes basal teleosts and primitive extant teleosts such as elopomorphs, osteoglossomorphs, and salmonids. The diural caudal skeleton is redefined as having two independent ural chordacentra plus their arcocentra, or two ural chordacentra plus their autocentra and arococentra, or only two ural chordacentra. A polyural caudal skeleton is identified by more than two ural centra, variably formed as given for the diural condition. The two ural centra of primitive teleosts may result from early fusion of ural centra 1 and 2 and of ural centra 3 and 4, or 3, 4, and 5 (e.g., elopomorphs), respectively. The two centra may corespond to ural centrum 2 and 4 only (e.g., salmonids). Additionally, ural centra 1 and 3 may be lost during the evolution of teleosts. Additional ural centra form late in ontogeny in advanced salmonids, resulting in a secondary polyural caudal skeleton. The hypural, which is a haemal spine of a ural centrum, results by growth and ossification of a single basiventral ural arococentrum and its haemal spine. The proximal part of the hypural always includes part of the ventral ural arcocentrum. The uroneural is a modification of a ural neural arch, which is demonstrated by a cartilaginous precursor. The stegural of salmonids and esocids originates from only one paired cartilaginous dorsal arcocentrum that grows anteriorly by a perichondral basal ossification and an anterodorsal membranous ossification. The true epurals of teleosts are detached neural spines of preural and ural neural arches as shown by developmental series; they are homologous to the neural spines of anterior vertebrae. Free epurals without any indication of connection with the dorsal arococentra are considered herein as an advanced state of the epural. Caudal distal radials originate from the cartilaginous distal portion of neural and haemal spines of preural and ural (epurals and hypurals) vertebrae. Therefore, they result from distal growth of the cartilaginous spines and hypurals. Cartilaginous plates that support rays are the result of modifications of the plates of connective tissue at the posterior end of hypurals (e.g., between hypurals 2 and 3 in salmonids) and first preural haemal spines, or from the distal growth of cartilaginous spines (e.g., epural plates in Thymallus). Among salmonids, conditions of the caudal skeleton such as the progressive loss of cartilaginous portions of the arcocentra, the progressive fusion between the perichondral ossification of arcocentra and autocentra, the broadening of the neural spines, the enlargement and interdigitation of the stegural, and other features provide evidence that Prosopium and Thymallus are the most primitive, and that Oncorhynchus and Salmo are the most advanced salmonids respectively. This interpretation supports the current hypothesis of phylogenetic relationships of salmonids. © 1992 Wiley-Liss, Inc.     

Development of the vertebral column and caudal complex in a flyingfish,Parexocoetus mento mento (Teleostei: Exocoetidae)

The developmental pattern of the vertebral column and caudal complex in juvenile (16.9 mm SL) to adult (112.2 mm SL)Parexocoetus mento mento is described Juvenile external caudal morphology was similar to the adult condition, although juveniles exhibited various internal ontogenetic changes. Osteological develoment was almost completed at 60–69 mm SL. Complete ossification of the vertebral column and caudal complex appeared to be the optimal condition giving strength for flight. Loss of perforations in the centra, neural and haemal arches may be consistent with the rigid and straightened body position during take-off. Some ontogenetic changes in the caudal complex were related to functional aspects. Ankylosis of the NPU2 spur to the uroneural notch, fusion of hypurals 3+4 and 5 and the elongated hypural 1+2 (lower hypural) were linked to the acquisition of stability and strength in the caudal complex.     

The composition of the caudal skeleton of teleosts (Actinopterygil: Osteichthyes)

The diural caudal skeleton of teleostean actinopterygians develops phylogeneticaily and ontogenetically from a polyural skeleton. The reduction of the polyural anlage to four, three, two or fewer centra in the adult caudal skeleton takes different pathways in different genera (e.g. compare Elops and Albula) and groups of teleosts. As a result, ural centra are not homologous throughout the teleosts. By numbering the ural centra in a homocercal tail in polyural fashion, one can demonstrate these and the following differences. The ventral elements (hypurals) always occur in sequential series, whereas the dorsal elements (epurals and uroneurals) may alter like the ural centra. The number of epurals, five or four in fossil primitive teleosts, is reduced in other primitive and advanced teleosts, but the same epurals are not always lost. The number of uroneurals, seven in fossil teleosts, is reduced in living teleosts, but it has not been demonstrated that the first uroneural is always derived from the neural arch of the same ural centrum. The landmark in the homocercal tail is the preural centrum I which can be identified by (1) bifurcation of the caudal artery and vein in its ventral element, the parhypural, (2) its position directly caudal to the preural centrum (PU2) which supports the lowermost principal caudal ray with its haemal spine, (3) carrying the third hypaxial element ventral to the course of arteria and vena pinnalis, and (4) by carrying the first haemal spine (parhypural) below the dorsal end of the ventral cartilage plate. The study of the development of the vertebral column reveals that teleosts have different patterns of centrum formation. A vertebral centrum is a complete or partial ring of mineralized, cartilaginous or bony material surrounding at least the lateral sides of the notochord. A vertebral centrum may be formed by arcocentrum alone, or arcocentral arcualia and chordacentrum, or arco-, chorda- and autocentrum, or arcocentral arcualia and autocentrum. This preliminary research demonstrates that a detailed ontogenetic interpretation of the vertebral centra and of the caudal skeleton of different teleosts may be useful tools for further interpretations of teleostean interrelationships.     

The caudal skeleton of ostariophysan fishes (Teleostei): Intraspecific variation in trichomycteridae (Siluriformes)

The different elements of the caudal skeleton of the South American catfish genera Nematogenys (Nematogenyinae) and Trichomycterus, Hatcheria, and Bullockia (Pygidiinae) (Siluriformes, Trichomycteridae) show Ontogenetic transformation of the second ural centrum in Trichomycteridae separates the subfamilies Nematogenyinae and Pygidiinae. In the former, the second ural centrum is aligned with the first ural centrum in early stages of ontogeny; it is not fused with the bases of hypurals 3 and 4 in any stage of development. In the Pygidiinae, in contrast, the second ural centrum is connected with the base of hypural 3 from an early stage of development on. One of the most noteworthy features of the Pygidiinae is the epural, a polymorphic element with three or four morphotypes that are species specific. The primitive catfish Nematogenys shows intraspecific variation in the ural centra, segmentation of procurrent caudal rays, and principal caudal ray formulae. Species of Trichomycterus, Hatcheria, and Bullockia are characterized by great intraspecific variability that involves ural centra, the epural, hypurapophyses, and the neural arches of the compound centrum. There is intraspecific variation in the fusion of the hypurals in some species of Trichomycterus. Intraspecific variation of the caudal skeleton of fishes of the family Trichomycteridae involves the presence and frequency of different morphotypes of the epural, neural arch of the compound centrum, fusion of hypurals, and principal caudal ray formulae. Ontogenetic changes of the first and second ural centra, hypurapophyses (with the exception of Nematogenys), and segmentation of procurrent caudal rays (in Nematogenys) are involved also.     

Leis' conundrum: homology of the clavus of the ocean sunfishes. 1. Ontogeny of the median fins and axial skeleton of Monotrete leiurus (Teleostei, Tetraodontiformes, Tetraodontidae)

We describe the ontogeny of the axial skeleton and median fins of the Southeast Asian freshwater puffer Monotrete leiurus, based on a reared developmental series. Most elements of the axial skeleton in M. leiurus arise in membrane bone. Only the base of the anterior three neural arches, the base of the hemal arches of the third preural centrum, the neural and hemal arches and spines of the second preural centrum, the parhypural, the two hypural plates, and the single epural are preformed in cartilage. In contrast to most teleosts, the proximal-middle radials of the dorsal and anal fins are upright and symmetrical and their distal tips coalesce during development to form a deep band of cartilage, from which the spherical distal radials are spatially separated.     

Reevaluation of the caudal skeleton of some actinopterygian fishes: II. Hiodon,Elops, and Albula

The vertebral centra of Hiodon, Elops, and Albula are direct perichordal ossifications (autocentra) which enclose the arcocentra as in Amia. An inner ring of ovoid cells forms in late ontogeny from the intervertebral space inside the autocentrum. The chordacentrum is reduced or completely absent in centra of adult Elops, whereas it forms an important portion of the centra in adult Hiodon. The posterior portion of the compound ural centrum 3+4+5 is partially (Hiodon) or fully formed by the chordacentrum (Elops, Albula). The haemal arches and hypurals are fused medially by cartilage or bone trabecles of the arcocentrum with the centra, even though they appear autogenous in lateral view in Elops and Albula. The composition of the caudal skeleton of fossil teleosts and the ontogeny of that of Hiodon, Elops, and Albula corroborate a one-to-one relationship of ural centra with these dorsal and ventral elements. The first epural (epural 1) of Elops relates to ural centrum 1, whereas the first epural (epural 2) of Hiodon and Albula relates to ural centrum 2. In Albula, the first ural centrum is formed by ural centrum 2 only. With 4 uroneurals Hiodon has the highest number within recent teleosts. Juvenile specimens of Hiodon have eight, the highest number of hypurals within recent teleosts; this is the primitive condition by comparison with other teleosts and pholidophorids. Reduction of elements in the caudal skeleton is an advanced feature as seen within elopomorphs from Elops to Albula. Such reductions and fusions occur in osteoglossomorphs also, but the lack of epurals and uroneurals separates most osteoglossomorphs (except Hiodon) from all other teleosts.     

Ontogeny,variation, and homology in Salvelinus alpinus caudal skeleton (Teleostei: Salmonidae)

The ontogeny of the caudal skeleton in the Arctic charr, Salvelinus alpinus was examined using an extensive series of cleared and stained specimens. We demonstrate the presence of skeletal components never reported previously within the Salmonidae. In contrast to the generalized condition for salmonids, seven hypurals (instead of six), and four uroneurals (instead of three) have been found in some specimens. Variation in the number and condition of epurals is documented. New hypotheses are proposed concerning (1) relationships among centra and their associated elements, (2) phylogenetic distribution of caudal characters within the Salmonidae, and (3) homology of caudal components. Using the published phylogenetic hypotheses, we provide evidence, that a seventh hypural and a fourth uroneural are taxic atavism in salmonids. The development of the salmonid homocercal fin is discussed in the light of a polyural scheme based on evidences of a one‐to‐one relationship among ural centra and their associated elements. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Description of Careproctus patagonicus sp. nov. and C. magellanicus sp. nov. (Pisces: Scorpaeniformes) from the lower slope of Drake Passage

Two new species of Liparidae are described from Drake Passage (55°32·8' S, 65°54·3' W). Careproctus patagonicus differs from all other congeneric species in the following combination of characters: C 9 (1+4/4), epural and parhypural unfused to single hypural; gill slit above P base; lower pectoral lobe well developed; pectoral girdle with one ventral radial (0+0+0+1) with a central foramen; da 2·4, aAf 10·2% LS; disk 41·5% HL; some crater-like pits on posterior half of body; head and anterior part of body white. Careproctus magellanicus has the following diagnostic characters: A 46; P 24, well-developed lower pectoral lobe; scattered crater-like spots and some thumb-tack prickles on body; one suprabranchial pore; basal plate of the pectoral girdle with two fenestrae; four radials (3+1), the first with a ventral notch; HL 17·1, preD 18·5% LS. The structure of the endochondral pectoral girdle of both species is commented.     

Redescription of a rare bothid,Asterorhombus bleekeri (Macleay), and description of a new species ofAsterorhombus from northwestern Australia (Teleostei: Pleuronectiformes)

A rare Australian bothid flounder.Asterorhombus bleekeri (Macleay), is redescribed from the holotype and ten additional specimens from the east coast of Qeensland, Gulf of Carpentaria and Rowly Shoal (Western Australia). The species is transferred fromArnoglossus Bleeker toAsterorhombus Tanaka because of the lack of obvious sexual dimorphism in the interorbital width and pectoral fin length, the lack of rostral and orbital spines, the yellow-white blind side body coloration, and the deeply cleft parhypural and hypural plates. The definition ofAsterorhombus was emended as follows: the first dorsal fin ray continuous with or separated from remaining fin rays and gill rakers slender or stubby, with or without serrations.Asterorhombus osculus sp. nov., formerly briefly described in the literature as unidentified species ofEngyprosopon, was described from eight specimens from the northwestern coast of Australia. The new species is most similar toA. bleekeri in lacking sexual dimorphism, and having the caudal skeleton with deep clefts, two or three rows of teeth on the upper jaw and a pair of conspicuous black spots on the caudal fin, in addition to a similar general appearance, but is distinguished from the latter by shorter gill rakers, a very small mouth and feebly ctenoid scales on the ocular side. Both species clearly differed fromA. intermedius andA. fijiensis in having two (or three) rows of teeth on the upper jaws, slender gill rakers without serrations, first dorsal fin ray continuous with the other fin rays, and a pair of conspicuous black spots on the caudal fin.     

Early development of fin-supports ani fin-rays in the milkfishChanos chanos

Development of fin-supports and fin-rays was observed in larval and juvenileChanos chanos, Chondrification of the caudal complex started at 4.70 mm SL. Ossification of the caudal elements started at 7.80 mm SL and was nearly completed at about 30 mm SL. Cartilaginous fusion of caudal elements, which occurs in hypurals of higher teleostean fishes but is not seen in lower teleosts, was observed between the neural arch of the preural centrum 1 and that of the ural centrum 1 via a small cartilage bridging the distal tips of the two arches. Caudal finrays began to develop at 6.60 mm SL, and an adult complement of principal rays was attained at 7.35 mm SL. Dorsal and anal pterygiophore elements were first evident at 6.70 mm and 6.65 mm SL, respectively. All proximal radiais were formed at 8.15 mm SL in both fins. Formation of dorsal and anal fin-rays started simultaneously at 8.60 mm SL, and adult fin-ray complements were attained at 10,00 mm and 10.70 mm SL, respectively. In the pectoral fin, the cleithrum, coraco-scapular cartilage and blade-like cartilage (fin plate) had already been formed at 4.65 mm SL. The mesocoracoid was observed to originate from the coraco-scapular cartilage and become detached from it in the course of ossification. Pectoral fin-ray formation started at 13.80 mm SL and was completed in number of rays at 20.00 mm SL. In the pelvic fin, the basipterygium was first evident at 13.00 mm SL. Pelvic fin-rays appeared at 13.80 mm SL and attained their adult count at 17.15 mm SL.     

When two equals three: developmental osteology and homology of the caudal skeleton in carangid fishes (Perciformes: Carangidae)

Ontogeny often provides the most compelling evidence for primary homology in evolutionary developmental studies and is critical to interpreting complex structures in a phylogenetic context. As an example of this, we document the ontogenetic development of the caudal skeleton of Caranx crysos by examining a series of cleared and stained larval and postlarval specimens. By studying ontogeny, we are able to more accurately identify some elements of the adult caudal skeleton than is possible when studying the adult stage alone. The presence of two epurals has been used as a synapomorphy of Caranginae (homoplastically present in the scomberoidine genera Scomberoides and Oligoplites). Here we find that three epurals (ep) are present in larvae and small postlarval juveniles (i.e.,<25 mm standard length [SL]) of C. crysos and other carangines, but ep2 never ossifies and does not develop beyond its initial presence. Ep2 was last observed in a 33.6 mm SL specimen as a small nodule of very lightly stained cartilage cells and eventually disappears completely. Therefore, the two epurals present in the adult are ep1 and ep3. In other carangines examined (e.g., Selene, Selar), the rudimentary ep2 ossifies and appears to fuse to the proximal tip of ep1. In these taxa, therefore, the two epurals of the adult appear to be ep1+2 and ep3. We found no indication of three epurals at any stage in the development of Oligoplites (developmental material of Scomberoides was unavailable). We discuss the osteology of the caudal skeleton of carangoid fishes generally and emphasize the power and importance of ontogeny in the identification of primary homology.     

Senegalese sole bone tissue originated from chondral ossification is more sensitive than dermal bone to high vitamin A content in enriched Artemia

Several studies have evaluated the effects of dietary vitamin A (VA) on the incidence of skeletal deformities during early ontogeny of fish, but little is known about its effects on bones depending on their process of ossification (dermal or chondral). We examined the incidence of skeletal deformities along development (30 and 48 dph) by double staining technique, in dermal (haemal and caudal vertebral bodies) and chondral (neural and haemal spines, epural, parahypural and hypurals) bones in Senegal sole post metamorphosed larvae fed with different dietary VA levels (37 000, 44 666, 82 666 and 203 000 UI total VA kg^?1 DW) during Artemia feeding phase (6–37 dph, at 18°C). Results obtained in this study showed that dietary VA disrupted the skeletogenesis in Senegalese sole post metamorphosed larvae by increasing the incidence of skeletal deformities in the axial skeleton and caudal fin complex, which were dependent on both bone morphogenesis and ossification processes. Fish fed with the highest dietary VA content showed the highest incidence of skeletal deformities and its value increased along ontogeny. However, when we compared the incidence of deformities in skeletal structures considering their ossification process, most skeletal structures derived from chondral ossification showed a significant higher increase in deformity incidences in fish fed an excess of VA (44 666, 82 666 and 203 000 UI kg^?1 DW), however within chondral bones, hypurals deformity incidence only increased in sole larvae fed Artemia highest VA content. In contrast, this dietary dose‐response effect was only noted in dermal bones from fish fed the highest dose of VA (203 000 UI kg^?1 DW). In addition, the incidence of deformities in chondral bones increased even when the dietary imbalance of VA was corrected, whereas dermal bones were not affected at later ages. These results indicated that depending on the ossification process from which different skeletal structures are derived, bones might be differentially affected by high dietary VA content. Those directly originated from the connective tissue with a preliminary cartilage stage were more sensitive to dietary VA excess than those formed by intramembranous ossification.     

Osteological development of the lophiid anglerfish,lophius gastrophysus

The osteological development of the head skeleton and dorsal, pectoral, and anal fin supports, are described from cleared and stained specimens ofLophius gastrophysus larvae, ranging from 4.6 to 21.8 mm NL; the results are compared with those of juvenile (79.8 mm SL) and adult (398 mm SL) specimens. Tiny conical teeth are present on the premaxillary, dentary, palatine and vomer since early stage. The first three dorsal fin spines are initially positioned on the midline of body posterior to the supraoccipital, but they migrate forward with growth and become cephalic in juveniles. The forward movement of the dorsal spines is produced by the forward extension of the cartilaginous basal inside the subepidermal space. During the planktonic larval stage the pectoral fins are on the sides of body as in ordinary fishes, but they move ventrad and become leg-like in bottom living juveniles and adults. Ossification of the caudal complex ofL. gastrophysus larvae proceeds very slowly and only the 21.8 mm NL larva has an almost completely ossified caudal complex. Eight principal caudal rays are loosely attached on the posterior edge of the hypurals and no procurrent rays are present. Larvae have well developed parhypurapophysis at the mid-portion of the urostyle which transforms into keel-like structure in juveniles and adults.     

Early scale development in Heterodontus (Heterodontiformes; Chondrichthyes): a novel chondrichthyan scale pattern

In two species of Heterodontus, H. portusjacksoni and H. galeatus, the first scales to develop form two opposing rows along the caudal fin axis on both the left and right sides of the fin. The opposing rows originate from an initial scale located on either side of the posterior tip of the caudal fin, with subsequent scales erupting in a posterior to anterior direction along the tail axis. These scale rows may strengthen tail movements, providing aeration in the egg case, but are lost later in ontogeny. Development of subsequent body scales shows a more irregular origin and arrangement, from anterior to posterior, to cover the dorsal and ventral lobes of the caudal fin. Although the early developmental pattern of the scale associated with the Heterodontus caudal fin has not been previously described, several chondrichthyan taxa, including chimeroids, likewise possess ordered rows of flank scales early in ontogeny that are subsequently lost. These ordered scales contrast with previous suggestions that chondrichthyan scale development is entirely random. Instead, regulated and sequential development of scales may be a plesiomorphic character for both chondrichthyans and osteichthyans, with the less organized arrangement in later ontogenetic stages being a derived condition within Chondrichthyes.     

Functional morphology of the caudal skeleton in teleostean fishes

The basic function of the caudal skeleton in teleostean fishes is to support the caudal fin, but its parts contribute to this function in somewhat different ways. The main axis for this support is the upturned terminal end of the vertebral column, which ends at the base of the uppermost principal rays. The uroneural struts just ahead of this axis provide support for it. The parts of the caudal skeleton behind and below this upturned axis, the hypurals and parhypural, not only support the caudal rays but also provide a means for differential movements between the upper and lower parts of the fin base. This basic caudal skeleton varies with the position of the fish in the sequence of teleosten evolution, the way in which the fish uses its caudal fin, and to some extent with the shape of the fin.     

The hemiramphid,Oxyporhamphus, is a flyingfish (exocoetidae)

Osteological and myological studies on the caudal complex of flying fishes (Exocoetidae) plusOxyporhamphus (of Hemiramphidae) revealed the following shared derived conditions: 1) neural spines of preural vertebrae broader than haemal spines; 2) spur present on posterior margin of preural vertebra 2;3) upper hypural plates steeply angled; 4) lower hypural plate extending strongly posteriorly; 5) lower hypural plate deeply surrounded by caudal fin rays; 6) flexor ventralis well developed, arising from neural spines; 7) flexor ventralis externus well developed; 8) adductor dorsalis well developed.Oxyporhamphus and exocoetids also share a lower jaw of adults not elongate and the premaxilla with a straight anterior margin. Therefore,Oxyporhamphus is transferred to the Exocoetidae, with which it shares a total of 10 derived conditions.     

Cranial and appendicular ontogeny of Bactrosaurus johnsoni,a hadrosauroid dinosaur from the Late Cretaceous of northern China

Abstract: The juvenile anatomy of various cranial and appendicular elements of the hadrosauroid dinosaur Bactrosaurus johnsoni is described in detail. Growth changes are documented from juvenile to adult stages for each skeletal element available. In the studied skull, ontogenetic trends consist of: development of features on the ventral surface of the frontal; reduction in the slope of the posteromedial process of the premaxilla; a posterior shift of the dorsal process of the maxilla; development of concavities on the medial surface of the prefrontal; increased robustness and development of the ventral flange of the jugal; decreased curvature of the long axis of the quadrate; increased ventral deflection of the dentary; and changes in the length/width proportions and depth of the anterior surface of the predentary. In the appendicular skeleton, the majority of ontogenetic variation from juvenile to adult occurs in the limb bones, including increased robustness of the deltopectoral crest of the humerus; relative shortening of the ulna; increased development of the fourth trochanter and mediolateral widening of the distal end of the femur; increased expansion of the cnemial crest of the tibia; and the increased prominence of articular protuberances and flanges of the metatarsals. A survey of the phylogenetically informative characters present in B. johnsoni indicates that several characters concerning the frontal, maxilla, jugal, quadrate, predentary, dentary, scapula, humerus and ilium are affected by ontogeny. Nevertheless, the majority of phylogenetic characters are not ontogenetically variable, suggesting that a substantial amount of the information provided by juvenile and subadult specimens for phylogenetic inference is reliable in basal hadrosauroids.     

An embryological study of ventralization of dorsal structures in the tail of medaka (Oryzias latipes) Da mutants

In adult Da ( double anal fin ) mutants of medaka ( Oryzias latipes ), structures such as the dorsal fin and the dorsal half of the caudal fin are ventralized in adult fish. However, there have been few embryological studies of the development of mutant phenotypes except those of the caudal fin. In this study, development of mutant phenotypes of the tail where they typically develop was examined morphologically at various stages of embryogenesis. The arrangement of melanocytes along the dorsal midline, the shape of the dorsal fin fold, and the shape of the dorsal myotome exhibited a ventral pattern in the tail at various embryonic stages in Da mutants.