Lessons from the first dorsal fin in atheriniforms—A new mode of dorsal fin development and its phylogenetic implications

The median fins in extant actinopterygians are the product of millions of years of evolution. During this time, different developmental patterns for the dorsal and anal fins emerged leading to a high variation in median fin morphology and ontogeny. In this study, the development of anal and dorsal fins in atheriniforms is described and its consequences for the current phylogenetic hypothesis are discussed. Developmental series of five atheriniform species were investigated using clearing and staining as well as antibody staining. The skeletal elements of the second dorsal fin and the anal fin emerge in a bidirectional pattern. The first dorsal fin, however, arises separately in front of the second dorsal fin after this one is almost completely formed. The pterygiophores of the first dorsal fin, including the interdorsal pterygiophores, develop from caudal to rostral, but the fin‐spines of the first dorsal fin form in the opposite direction. This new mode of fin development has been found in all examined atheriniform species with two dorsal fins. Several morphological characters of atheriniforms, including interdorsal pterygiophores, are also found in one other taxon: the Mugiliformes. Thus, several dorsal fin characteristics may provide evidence for a closer relationship of these two taxa.     

The dorsal fin engine of the seahorse (Hippocampus sp.)

The muscles, fin ray joints, and supporting structures underlying the dorsal fin are described for two seahorse species: Hippocampus zosterae and Hippocampus erectus. A fan-shaped array of cartilaginous bones, the pterigiophores, form the internal supporting structure of the dorsal fin. Each pterigiophore is composed of a proximal radial that extends from a vertebra to the dorsal side of the animal, where it fuses to a middle radial. The middle radials fuse with each other to form a dorsal ridge upon which sit the spheroidal distal radials. Each distal radial articulates with a fin ray on its dorsal side and is attached to the dorsal ridge on its ventral side by a material that has been histologically identified as elastic cartilage. Together these connections form a two-axis joint that permits elevation, depression, and inclination of the ray. Each fin ray is actuated by two bilateral pairs of muscles, an anterior pair of inclinators, and a posterior pair of depressors. The anteriormost fin ray is actuated by three bilateral pair of muscles, the inclinators, the depressors, and a pair of elevator muscles that are positioned anterior to the inclinators. Preliminary examinations of the ray joints of the pectoral and anal fins of adult H. zostera and the pectoral fins of newborn H. erectus revealed structures similar to that seen in the dorsal fins. To further explore the structure and function of the dorsal fin gross dissections and simple functional tests were performed on H. erectus and H. barbouri and behavioral observations were made of all three species plus Hippocampus kuda.     

Morphological Development of the Opercular Apparatus in Kneria wittei(Ostariophysi: Gonorynchiformes) with Comments on its Possible Function

The structure, development and function of the male opercular apparatus were examined in the freshwater ostariophysan Kneria wittei using a combination of techniques (i.e. histology, SEM, skeletal clearing and staining). Specimens ranged in standard length from 10 to 65 mm. Morphological comparisons were also made between males and females, as well as comparisons with closely related taxa. Our study shows that the Kneria opercular apparatus consists of two structures: an opercular cup and a posteriorly positioned opercular flange, each of which is composed largely of connective tissue with keratinized papillae and ridges positioned on the surface. Our results further show that the development of the opercular apparatus is linked directly to the development of the anterior vertebral elements (i.e. epicentral intermuscular bones and neural arches). Understanding the function of the opercular apparatus must therefore include an understanding of its associated skeletal elements. As a result of our morphological observations two hypotheses for the function of the opercular unit are addressed; as an auditory specialization, and as an adhesive device. Copyright     

Musculoskeletal morphology and regionalization within the dorsal and anal fins of bluegill sunfish (Lepomis macrochirus)

Ray‐finned fishes actively control the shape and orientation of their fins to either generate or resist hydrodynamic forces. Because of the emergent mechanical properties of their segmented, bilaminar fin rays (lepidotrichia), and actuation by multiple muscles, fish can control the rigidity and curvature of individual rays independently, thereby varying the resultant forces across the fin surfaces. Expecting that differences in fin‐ray morphology should reflect variation in their mechanical properties, we measured several musculoskeletal features of individual spines and rays of the dorsal and anal fins of bluegill sunfish, Lepomis macrochirus, and assessed their mobility and flexibility. We separated the fin‐rays into four groups based on the fin (dorsal or anal) or fin‐ray type (spine or ray) and measured the length of the spines/rays and the mass of the three median fin‐ray muscles: the inclinators, erectors and depressors. Within the two ray groups, we measured the portion of the rays that were segmented vs. unsegmented and branched vs. unbranched. For the majority of variables tested, we found that variations between fin‐rays within each group were significantly related to position within the fin and these patterns were conserved between the dorsal and anal rays. Based on positional variations in fin‐ray and muscle parameters, we suggest that anterior and posterior regions of each fin perform different functions when interacting with the surrounding fluid. Specifically, we suggest that the stiffer anterior rays of the soft dorsal and anal fins maintain stability and keep the flow across the fins steady. The posterior rays, which are more flexible with a greater range of motion, fine‐tune their stiffness and orientation, directing the resultant flow to generate lateral and some thrust forces, thus acting as an accessory caudal fin. J. Morphol., 2012. © 2011 Wiley Periodicals, Inc.     

A specimen of Salmo trutta (Pisces: Salmonidae) possessing an additional dorsal fin

A specimen of brown trout, Salmo trutta , with a supernumerary dorsal fin located just posterior to the nape is documented from the Yarra River, Victoria, Australia.     

Structure of supporting elements in the dorsal fin of percid fishes

The dorsal fin is one of the most varied swimming structures in Acanthomorpha, the spiny‐finned fishes. This fin can be present as a single contiguous structure supported by bony spines and soft lepidotrichia, or it may be divided into an anterior, spiny dorsal fin and a posterior, soft dorsal fin. The freshwater fish family Percidae exhibits especially great variation in dorsal fin spacing, including fishes with separated fins of varying gap length and fishes with contiguous fins. We hypothesized that fishes with separated dorsal fins, especially those with large gaps between fins, would have stiffened fin elements at the leading edge of the soft dorsal fin to resist hydrodynamic loading during locomotion. For 10 percid species, we measured the spacing between dorsal fins and calculated the second moment of area of selected spines and lepidotrichia from museum specimens. There was no significant relationship between the spacing between dorsal fins and the second moment of area of the leading edge of the soft dorsal fin.     

The morphology of the club glands on the dorsal fin of mature male shannies, Lipophvys pholis (L.) (Blenniidae: Teleostei)

The pelvic, pectoral, anal and dorsal fins of mature male, female and immature male Lipophrys pholis (L.) were examined by light microscopy for the presence of club glands. All fins except the dorsal showed a highly stratified epidermis and a thick cuticle. Club glands were present on the dorsal fin of mature male fisH but only during the breeding season. The development and decline of the club glands corresponds to the period of gonadal build-up and spawnout. Each club gland comprises several thousand bundles ofcolumnar cells. The columnar cells surround acentral pore which opens to the outside through a layer of Malpighian cells. The substance produced by the gland includes mucopolysacchdride. The function of the secretion is unknown and is discussed in relation to studies on similar glands in several Mediterranean blenniids.     

The biogeography of otophysan fishes (Ostariophysi: Otophysi): a new appraisal

Aim To present a new hypothetical history of the otophysan fishes. Location World‐wide. Methods Utilization of recent information about otophysan phylogeny, palaeontology, biogeography and external relations. Results Anatomical, mitochondrial DNA and karological research has indicated that the Cypriniformes is the most primitive of the four orders that comprise the Otophysi. The Siluriformes is next in the phylogenetic progression followed by the Characiformes and the Gymnotiformes. A molecular clock estimate indicated that the divergence time between the cypriniforms and the remaining otophysans took place about 250 Ma. But, otophysans have apparently never occurred on Madagascar‐India and that land mass did not separate from Africa until 158–160 Ma. As we know that some otophysans existed in Africa–South America prior to 115 Ma, they must have originated between 115 and 160 Ma. Main conclusions Comparison of phylogeny and biogeography appears to indicate that the Otophysi originated in South America, the location of the apomorphic Gymnotiformes. It seems that the gymnotiforms did not radiate in time to get across to Africa and the characiforms did not expand in time to catch the connection to Asia. Apparently, it was only the ancestors to the cypriniform and siluriform lines that expanded early enough to reach Asia in the late Jurassic. There remains a gap of about 80 Myr between the oldest known otophysan fossil and the approximate time of dispersal to Asia. Further progress awaits more phylogenetic or fossil discoveries.     

Development of the pectoral, pelvic, dorsal and anal fins in cultured sea bream

The pectoral fin girdle was the first element of the fins to develop in Sparus aurata. By 3·1mm L _N (notochord length) the cleithrum was ossified and the cartilaginous caracoid-scapula was present. The fin was fully developed at 11·6 mm L _S (standard length) and by 16·0 mm L _S most elements of the fin were ossified. The pelvic fins were the last pair to develop and rudiments of these were first detected at 7·9 mm L _S. The pelvic fin and girdle were completely formed and ossified at 16·0 mm L _S. The development of dorsal and anal fins began at c. 6·5–7·0 mm L _S with the formation of 10 cartilaginous dorsal proximal radials and eight cartilaginous ventral proximal radials. The three cartilaginous predorsals (supraneurals) appeared at 7·7 mm L _S and the ossification of dorsal and anal proximal and distal radials began, respectively, at 10·5 mm L _S and 11·3 mm L _S. Ossified structures in the fins were also classified according to their origin, as being either dermal or endochondral. Finally the chronology of appearance of fin structures in S. aurata was compared with that reported for other Sparidae, Engraulidae and Haemulidae.     

Differentiation of chondrocytes and scleroblasts during dorsal fin skeletogenesis in flounder larvae

In teleosts, the embryonic fin fold consists of a peridermis, an underlying epidermis and a small number of mesenchymal cells. Beginning from such a simple structure, the fin skeletons, including the proximal and distal radials and lepidotrichia (finrays), develop in the dorsal fin fold at the larval stage. Their process of skeletogenesis and embryonic origin are unclear. Using flounder larvae, we report the differentiation process for chondrocytes and scleroblasts prior to fin skeletogenesis and the effects of retinoic acid (RA) on it. In early larvae, the mesenchymal cells grow between the epidermis and spinal cord to form a line of periodical condensations, which are proximal radial primordia, to produce chondrocytes. The prescleroblasts, which ossify the proximal radial cartilages, differentiate in the mesenchymal cells remaining between the cartilages. Then, mesenchymal condensations occur between the distal ends of the proximal radials, forming distal radial primordia, to produce chondrocytes. Simultaneously, condensations occur between the distal radial primordia and peridermis, which are lepidotrichia primordia, to produce prescleroblasts. Exogenous RA specifically inhibits the mesenchymal condensation prior to the proximal radial formation together with the down-regulation of sonic hedgehog (shh) and patched (pta) expression, resulting in the loss of proximal radials. Thus, it was indicated that differentiation of the precursor cells of radials and lepidotrichia begins in the proximal part of the fin fold and that the initial mesenchymal condensation prior to the proximal radial formation is highly susceptible to the effects of RA. Lepidotrichia formation does not occur where proximal radials are absent, indicating that lepidotrichia differentiation requires interaction with the radial cartilages. To examine the suggestion that neural crest cells contribute to the medial fin skeletons, we localized the HNK-1 positive cells in flounder embryos and slug and msxb-positive cells in pufferfish, Fugu rubripes, embryos. That the positive cells commonly arrive at the proximal part of the fin fold does not contradict the suggestion, but their final destiny as radial chondrocytes or lepidotrichia scleroblasts, should be further investigated.     

Microanatomy of the paired‐fin pads of ostariophysan fishes (Teleostei: Ostariophysi)

Members of the teleost superorder Ostariophysi dominate freshwater habitats on all continents except Antarctica and Australia. Obligate benthic and rheophilic taxa from four different orders of the Ostariophysi (Gonorynchiformes, Cypriniformes, Characiformes, and Siluriformes) frequently exhibit thickened pads of skin along the ventral surface of the anteriormost ray or rays of horizontally orientated paired (pectoral and pelvic) fins. Such paired‐fin pads, though convergent, are externally homogenous across ostariophysan groups (particularly nonsiluriform taxa) and have been considered previously to be the result of epidermal modification. Histological examination of the pectoral and/or pelvic fins of 44 species of ostariophysans (including members of the Gonorynchiforms, Cypriniformes, Characiformes, and Siluriformes) revealed a tremendous and previously unrecognized diversity in the cellular arrangement of the skin layers (epidermis and subdermis) contributing to the paired‐fin pads. Three types of paired‐fin pads (Types 1–3) are identified in nonsiluriform ostariophysan fishes, based on differences in the cellular arrangement of the epidermis and subdermis. The paired‐fin pads of siluriforms may or may not exhibit a deep series of ridges and grooves across the surface. Two distinct patterns of unculus producing cells are identified in the epidermis of the paired‐fin pads of siluriforms, one of which is characterized by distinct bands of keratinization throughout the epidermis and is described in Amphilius platychir (Amphiliidae) for the first time. General histological comparisons between the paired fins of benthic and rheophilic ostariophysan and nonostariophysan percomorph fishes are provided, and the possible function(s) of the paired‐fin pads of ostariophysan fish are discussed. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Wing design and morphology of the harbor porpoise dorsal fin

The correlation between skin structure and hydrodynamic design of the dorsal fin of the harbor porpoise (Phocoena phocoena) was examined. For the study of fin morphology and geometry, a scheme of sampling representing a two-parameter mesh on the fin surface was used. At each data point the thickness of the epidermis, papillary and subpapillary layers of the dermis, the ligamentous layer of the fin, as well as the angle formed by the direction of dermal ridges and the fin root chord were measured. On the basis of fin cross-sections the three-dimensional surface models of the fin in a 1 : 1 scale were created with a CAD program. The shape of the model was evaluated by the wing and hydrofoil parameters (angle of leading edge sweep, leading edge radius, maximum thickness of the fin cross-section, and position of maximum thickness from the leading edge). Hydrodynamic performance of the fin cross-sections was studied with a CFD program. Regional variability of the parameters of morphology was compared with spanwise variability of the parameters of cross-sectional geometry. It was found that skin structure parameters correlate with the hydrodynamically relevant parameters of the fin and fin cross-sections. Regularities of skin structure of the harbor porpoise dorsal fin are considered indirect evidence of the adaptation of porpoise skin to the fin flow.     

The anal fin complex in a weakly discharging electric fish, Gnathonemus petersii(Mormyridae)

An examination of the permanent bony structures of the anal fin complex in the mormyrid fish, Gnathonemus petersii , revealed two new structural sexual dimorphisms: longer proximal pterygiophores and wider anal fin rays in males than in females. Both structures are thought to facilitate the male's courtship‐associated anal fin reflex. Adult male mormyrid fishes are characterized by a dorsally directed indentation of the posterior body wall (anal fin indentation). The expression of this indentation in males, presumably driven by anal fin musculature, was correlated with the fish's gonadal state: large indentations were associated with high gonado‐somatic indices and small indentations with low indices.     

The median‐fin skeleton of the Eastern Atlantic and Mediterranean clingfishes Lepadogaster lepadogaster (Bonnaterre) and Gouania wildenowi (Risso) (Teleostei: Gobiesocidae)

Previous research on the osteology of the Gobiesocidae focused mostly on the neurocranium and the thoracic sucking disc (formed by the paired‐fin girdles). Little attention has been paid to the skeleton of the median fins. The dorsal‐ and anal‐fin skeleton of Lepadogaster lepadogaster and other gobiesocids (excluding Alabes, which lacks these fins) are characterized by the absence of spines, branched fin‐rays, and middle radials. In gobiesocids, the distal radials never ossify and consist of elastic hyaline‐cell cartilage. Gouania wildenowi is unique among gobiesocids in having further reductions of the dorsal‐ and anal‐fin skeleton, including a notable decrease in the size of the proximal‐middle radials in an anterior–posterior direction. Unlike L. lepadogaster, which exhibits a one‐to‐one relationship between the dorsal‐ and anal‐fin rays and proximal‐middle radials, G. wildenowi has a higher number of proximal‐middle radials than distal radial cartilages and fin rays in the dorsal and anal fins. In G. wildenowi, the dorsal‐ and anal‐fin rays do not articulate with the distal tip of the proximal‐middle radials but are instead positioned between proximal‐middle radials, which is unusual for teleosts. Previously unrecognized dorsal and ventral pads of elastic hyaline‐cell cartilage are also present in the caudal skeleton of L. lepadogaster, G. wildenowi, and all other gobiesocids examined. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

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.     

Anatomy and early development of the pectoral girdle,fin, and fin spine of sturgeons (Actinopterygii: Acipenseridae)

Acipenseriformes hold an important place in the evolutionary history of bony fishes. Given their phylogenetic position as extant basal Actinopterygii, it is generally held that a thorough understanding of their morphology will greatly contribute to the knowledge of the evolutionary history and the origin of diversity for the major osteichthyan clades. To this end, we examined comparative developmental series from the pectoral girdle in Acipenser fulvescens, A. medirostris, A. transmontanus, and Scaphirhynchus albus to document, describe, and compare ontogenetic and allometric differences in the pectoral girdle. We find, not surprisingly, broad congruence between taxa in the basic pattern of development of the dermal and chondral elements of the pectoral girdle. However, we also find clear differences in the details of structure and development among the species examined in the dermal elements, including the clavicle, cleithrum, supracleithrum, posttemporal, and pectoral‐fin spine. We also find differences in the internal fin elements such as the distal radials as well as in the number of fin rays and their association with the propterygium. Further, there are clear ontogenetic differences during development of the dermal and chondral elements in these species and allometric variation in the pectoral‐fin spine. The characters highlighted provide a suite of elements for further examination in studies of the phylogeny of sturgeons. Determining the distribution of these characters in other sturgeons may aid in further resolution of phylogenetic relationships, and these data highlight the role that ontogenetic and comparative developmental studies provide in systematics. J. Morphol. 276:241–260, 2015. © 2014 Wiley Periodicals, Inc.     

Development of osteological structures in the sea bream: vertebral column and caudal fin complex

The development of cartilaginous structures in cultured sea bream Sparus aurata larvae and the timing of their ossification was studied. In cultivated sea bream larvae the first cartilaginous structure to be identified was hypural 1 at 4.1 mm notochord length ( L _N). By 5.3 mm L _N, prior to the onset of ossification, it was possible to distinguish the following cartilaginous structures: all 23 neural arches, all 13 haemal arches and two of the four pairs of parapophyses. The neural arches 1–4 and 15–23 were formed on the notochord and elongated dorsally, while neural arches 5–14 appeared on the dorsal side of the spinal cord and elongated ventrally. Initiation of ossification occurred at 5.7–6.0 mm standard length ( L _S) when the cartilaginous ontogeny of the vertebral column was completed. Ossification was coincident with dorsal flexion at the posterior end of the notochord and occurred in a sequential manner: (1) dorsoanteriorly, the cartilaginous neural arches and the centra were the first structures to ossify; (2) ventrad at the centre, at 7.0–7.5 mm L _S; (3) posteriorly at 7.1 mm L _S the hypural complex and urostyle (24th centrum) were ossified; and (4) dorsad at the centre (neural arches and spines).     

The ontogeny and homology of the Weberian apparatus in the zebrafish Danio rerio (Ostariophysi: Cypriniformes)

The ontogeny of the Weberian apparatus was examined in the zebrafish, Danio rerio , using both cleared and stained specimens and histology. Over 300 individuals from four independent zebrafish lineages, ranging in size from 3 to 28 mm TL, were examined for this study. Results provide a basic understanding of the development of the Weberian apparatus in the wild-type zebrafish. Our results, in conjunction with those already published, point to substantial variation in the development of the Weberian apparatus among otophysans and new interpretations of the homology of certain ossicles (e.g. tripus and claustrum). Hypotheses of homology among various Weberian ossicles are considered and represent an important step in understanding the evolution of sound transmission in ostariophysan fishes.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 241–254.     

Paired fin skeletons and relationships of the fossil group Porolepiformes (Osteichthyes: Sardcopterygii)

The endoskeletal girdles, anocleithrum and paired fin supports of the porolepiform fish Glyptolepis (Osteichthyes: Sarcopterygii: Porolepiformes) are figured and described. The pectoral fin skeleton is known from the proximal part only and the pelvic fin skeleton is fragmentary, but the scapulocoracoid, anocleithrum and pelvic girdle can be reconstructed in their entirety. The anocleithrum is entirely subdermal. The pectoral fin skeleton in shown to be biserial, with a large number of axial mesomeres, whereas the pelvic fin contains fewer mesomeres and is strongly asymmetrical with very few postaxial radials. The scapulocoracoid is essentially similar to a reconstruction figured by Jarvik (1980), but has a more elongate glenoid; this has functional implications. The pelvic girdle consists of two separate halves as in Eusthenopteron, but differs from that genus in lacking dorsolateral rami. A brief survey of the evidence of paired fin structure in other porolepiform genera is carried out to establish whether the structures seen in Glyptolepis are likely to be representative for the Porolepiformes. A study of the morphology and muscle attachments of the paired fin skeletons indicates that the pattern of fin movement was significantly different from that in Neoceratodus. The fin supports and girdles of Glyptolepis are compared with those of other sarcopterygian groups as well as with actinopterygians, placoderms and sharks, in order to establish evolutionary polarities. Glyptolepis is shown to display a number of derived characters. The information gained from the comparison is used to construct a maximum parsimony cladogram, which places coelacanths as the sister group of porolepiforms + lungfishes, with the rhizodonts + tetrapods and osteolepiforms as successive sister groups of this clade. Characters of uncertain polarity are considered in the light of this cladogram. A comparison with recently published cladograms shows that none are completely compatible with the results from this study.