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.     

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.     

A comparison of pectoral fin ray morphology and its impact on fin ray flexural stiffness in labriform swimmers

The organization of tissues in appendages often affects their mechanical properties and function. In the fish family Labridae, swimming behavior is associated with pectoral fin flexural stiffness and morphology, where fins range on a continuum from stiff to relatively flexible fins. Across this diversity, pectoral fin flexural stiffness decreases exponentially along the length of any given fin ray, and ray stiffness decreases along the chord of the fin from the leading to trailing edge. In this study, we examine the morphological properties of fin rays, including the effective modulus in bending (E), second moment of area (I), segmentation, and branching patterns, and their impact on fin ray stiffness. We quantify intrinsic pectoral fin ray stiffness in similarly sized fins of two closely related species that employ fins of divergent mechanics, the flapping Gomphosus varius and the rowing Halichoeres bivittatus. While segmentation patterns and E were similar between species, measurements of I and the number of fin ray branch nodes were greater in G. varius than in H. bivittatus. A multiple regression model found that of these variables, I was always significantly correlated with fin ray flexural stiffness and that variation in I always explained the majority of the variation in flexural stiffness. Thus, while most of the morphological variables quantified in this study correlate with fin ray flexural stiffness, second moment of area is the greatest factor contributing to variation in flexural stiffness. Further, interspecific variation in fin ray branching pattern could be used as a means of tuning the effective stiffness of the fin webbing to differences in swimming behavior and hydrodynamics. The comparison of these results to other systems begins to unveil fundamental morphological features of biological beams and yields insight into the role of mechanical properties in fin deformation for aquatic locomotion.     

Variation in flexural stiffness of the lepidotrichia within and among the soft fins of yellow perch under different preservation techniques

Although the ray‐finned fishes are named for their bony, segmented lepidotrichia (fin rays), we are only beginning to understand the morphological and functional diversity of this key vertebrate structure. Fin rays support the fin web, and their material properties help define the function of the entire fin. Many earlier studies of fin ray morphology and function have focused on isolated rays, or on rays from only one or two fins. At the same time, relatively little is known about how different preservation techniques affect the material properties of many vertebrate structures, including fin rays. Here, we use three‐point bending tests to examine intra‐ and inter‐fin variation in the flexural stiffness of fin rays from yellow perch, Perca flavescens. We sampled fin rays from individuals that were assigned to one of three preservation treatments: fresh, frozen, and preserved with formalin. The flexural stiffness of the fin rays varied within and among fins. Pelvic‐fin rays were the stiffest, and pectoral fin rays the least stiff. The fin rays of the dorsal, anal, and caudal fins all had similar stiffness values, which were intermediate relative to those from the paired fins. The flexural stiffness of the fin rays was higher in rays that were at the leading edge of the fin. This variation in flexural stiffness was associated with variation in joint density and the relative length of the unsegmented proximal base of the fin rays. There was no significant difference in flexural stiffness between fresh and frozen specimens. In specimens preserved with formalin, there is a small but significant effect on stiffness in smaller fin rays.     

Early osteological development of the fins in the hatchery-reared red porgy, Pagrus pagrus (L. 1758)

The present study was undertaken to establish the normal, healthy features of morphological structures at various developmental stages as achieved under well-defined environmental culture conditions (temperature between 16 and 21°C, salinity 36 ppt, pH around 7.6, and oxygen saturation over 95%) common in aquaculture of the species. The pectoral fin supports began to develop at 2.90 mm total length (TL), followed by those of dorsal fins at 5.5 mm TL, caudal fins at 5.6 mm TL, pelvic fins at 5.9 mm TL and anal fins at 6.0 mm TL. The pelvic fins appeared fully at 7.4 mm TL. Development of dorsal lepidotrichia was first observed at 6.9 mm TL, attaining their final number at 7.6 mm TL. The dorsal spines first appeared at 6.5 mm TL and were complete at 7.4 mm TL. The anal lepidotrichia appeared during the development phase from 6.8 to 8.6 mm TL. At 5.6 mm TL, the upward flexion of the urostyle was initiated. The caudal lepidotrichia formed within the primordial fin at 5.6 mm TL and reached the final count at 7.4 mm TL. The caudal dermatotrichia first appeared at 7.3 mm TL and all forms were observed by 15.5 mm TL. The development pattern of fin supports found in Pagrus pagrus is quite similar to that described for other Sparid species.     

New genus and species of the family stichaeidae from Hokkaido,Japan

Specimens of a new genus and species of the stichaeid fish,Leptostichaeus pumilus, were collected from the Okhotsk Sea off Hokkaido in Japan. The present new genus and species clearly differs from all the other genera and species of the stichaeid fishes in the following characters: 3 or 4 pectoral fin rays; 10 or fewer caudal principal rays; 79–82 dorsal spines; no pelvic fin; last interneural spine supporting a single dorsal spine; infraorbital, occipital and lateral line canals absent; moderate size of dorsal spine shorter than eye diameter; membranes of dorsal and anal fins widely connected with caudal fin; a large black spot divided by a yellow band present just above gill cover.     

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.     

中国隆头鱼科一新纪录种——雀尖嘴鱼

尖嘴鱼属(Gomphosus)是一群分布于印度洋和太平洋热带珊瑚礁海域鱼类,共有2种,以往在中国海域记录有1种杂色尖嘴鱼(G.varius)。我们在分析20世纪90年代采自中国南海大陆坡的鱼类标本时,发现了该属的另一种雀尖嘴鱼(Gomphosus caeruleus Lacepède,1801),为中国新纪录种。本种的主要鉴别特征为:体呈浅黄褐色(雌)或深黑色(雄);吻部特别延长呈管状;体长为体高的4.2倍,为头长的2.6倍;背鳍Ⅷ-13,臀鳍Ⅲ-11,胸鳍i(不分支)+14(分支);脊椎骨25;鳃盖条7;体被中大圆鳞;侧线完全,在背鳍条的后部下方急剧向下弯折,侧线有孔鳞片27;头部仅鳃盖上部有9枚呈三角状排列的小鳞;背鳍起点前方有鳞8行;背鳍第一至第三鳍棘间的鳍膜具1黑斑;尾鳍截形。     

Neolumpenus unocellatus,a new genus and species of stichaeid fish from Japan

Neolumpenus unocellatus gen. et sp. nov., a stichaeid fish (subfamily Lumpeninae,sensu Makushok, 1958) is described on the basis of a single specimen found in the stomach of the Pacific cod,Gadus macrocephalus Tilesius, caught off Akkeshi, Hokkaido, Japan. The new genus and species is distinguished from all other lumpenines in having the following combination of characters: 1) 51 dorsal spines, 33 anal fin rays, 57 total vertebrae; 2) broad pelvic fin with deeply-branched soft rays; 3) lower rays of pectoral fin branched and not prolonged backward; 4) prevomerine and palatine teeth present; 5) pungent spines present in pelvic and anal fins; 6) upper lip fused to snout anteriorly; 7) gill openings not extending forward beyond a vertical through posterior margin of eye; 8) minimal (fifth) hypural present; 9) first interneural spine inserted between first and second neural spines; 10) extremely large cephalic sensory pores present; 11) high, steep snout; 12) ocellus on dorsal base of caudal fin.     

Correlated evolution of body and fin morphology in the cichlid fishes

Body and fin shapes are chief determinants of swimming performance in fishes. Different configurations of body and fin shapes can suit different locomotor specializations. The success of any configuration is dependent upon the hydrodynamic interactions between body and fins. Despite the importance of body–fin interactions for swimming, there are few data indicating whether body and fin configurations evolve in concert, or whether these structures vary independently. The cichlid fishes are a diverse family whose well‐studied phylogenetic relationships make them ideal for the study of macroevolution of ecomorphology. This study measured body, and caudal and median fin morphology from radiographs of 131 cichlid genera, using morphometrics and phylogenetic comparative methods to determine whether these traits exhibit correlated evolution. Partial least squares canonical analysis revealed that body, caudal fin, dorsal fin, and anal fin shapes all exhibited strong correlated evolution consistent with locomotor ecomorphology. Major patterns included the evolution of deep body profiles with long fins, suggestive of maneuvering specialization; and the evolution of narrow, elongate caudal peduncles with concave tails, a combination that characterizes economical cruisers. These results demonstrate that body shape evolution does not occur independently of other traits, but among a suite of other morphological changes that augment locomotor specialization.     

Actinotrichia collagens and their role in fin formation

The skeleton of zebrafish fins consists of lepidotrichia and actinotrichia. Actinotrichia are fibrils located at the tip of each lepidotrichia and play a morphogenetic role in fin formation. Actinotrichia are formed by collagens associated with non-collagen components. The non-collagen components of actinotrichia (actinodins) have been shown to play a critical role in fin to limb transition. The present study has focused on the collagens that form actinotrichia and their role in fin formation. We have found actinotrichia are formed by Collagen I plus a novel form of Collagen II, encoded by the col2a1b gene. This second copy of the collagen II gene is only found in fishes and is the only Collagen type II expressed in fins. Both col1a1a and col2a1b were found in actinotrichia forming cells. Significantly, they also expressed the lysyl hydroxylase 1 (lh1) gene, which encodes an enzyme involved in the post-translational processing of collagens. Morpholino knockdown in zebrafish embryos demonstrated that the two collagens and lh1 are essential for actinotrichia and fin fold morphogenesis. The col1a1 dominant mutant chihuahua showed aberrant phenotypes in both actinotrichia and lepidotrichia during fin development and regeneration. These pieces of evidences support that actinotrichia are composed of Collagens I and II, which are post-translationally processed by Lh1, and that the correct expression and assembling of these collagens is essential for fin formation. The unique collagen composition of actinotrichia may play a role in fin skeleton morphogenesis.     

Function of the dorsal fin in bluegill sunfish: Motor patterns during four distinct locomotor behaviors

The median fins of fishes are key features of locomotor morphology which function as complex control surfaces during a variety of behaviors. However, very few studies have experimentally assessed median fin function, as most workers focus on axial structures. In particular, the dorsal fin of many teleost fishes possesses both spiny anterior and soft posterior portions which may function separately during locomotion. We analyzed the function of the soft region of the dorsal fin and of the dorsal inclinator (Di) muscles which are the primary muscles responsible for lateral flexion. We used electromyography to measure in vivo Di activity, as well as activity of the red myomeric muscles located at a similar longitudinal position. We quantified motor patterns during four locomotor behaviors: braking and three propulsive behaviors (steady swimming, kick and glide swimming, and C-starts). During the three propulsive swimming behaviors, the timing of Di activity was more similar to that of ipsilateral red myomeric muscle rather than to contralateral myomeric activity, whereas during braking the timing of activity of the Di muscles was similar to that of the contralateral myomeric musculature. During the three propulsive behaviors, when the Di muscles had activity, it was consistent with the function of stiffening the soft dorsal fin to oppose its tendency to bend as a result of the body being swept laterally through the water. In contrast, activity of the Di muscles during braking was consistent with the function of actively flexing the soft dorsal fin towards the side of the fish that had Di activity. Activity of the Di muscles during steady speed swimming was generally sufficient to resist lateral bending of the soft dorsal fin, whereas during high speed kick and glide swimming and C-starts, Di activity was not sufficient to resist the bending caused by resistive forces imposed by the water. Cumulative data from all four behaviors suggest that the Di muscles can be activated independently relative to the myomeric musculature rather than having a single phase relationship with the myomeric muscle common to all of the observed behaviors. © 1996 Wiley-Liss, Inc.     

Experimental Hydrodynamics and Evolution: Function of Median Fins in Ray-finned Fishes

The median fins of fishes consist of the dorsal, anal, and caudal fins and have long been thought to play an important role in generating locomotor force during both steady swimming and maneuvering. But the orientations and magnitudes of these forces, the mechanisms by which they are generated, and how fish modulate median fin forces have remained largely unknown until the recent advent of Digital Particle Image Velocimetry (DPIV) which allows empirical analysis of force magnitude and direction. Experimental hydrodynamic studies of median fin function in fishes are of special utility when conducted in a comparative phylogenetic context, and we have examined fin function in four ray-finned fish clades (sturgeon, trout, sunfish, and mackerel) with the goal of testing classical hypotheses of fin function and evolution. In this paper we summarize two recent technical developments in DPIV methodology, and discuss key recent findings relevant to median fin function. High-resolution DPIV using a recursive local-correlation algorithm allows quantification of small vortices, while stereo-DPIV permits simultaneous measurement of x, y, and z flow velocity components within a single planar light sheet. Analyses of median fin wakes reveal that lateral forces are high relative to thrust force, and that mechanical performance of median fins (i.e., thrust as a proportion of total force) averages 0.35, a surprisingly low value. Large lateral forces which could arise as an unavoidable consequence of thrust generation using an undulatory propulsor may also enhance stability and maneuverability. Analysis of hydrodynamic function of the soft dorsal fin in bluegill sunfish shows that a thrust wake is generated that accounts for 12% of total thrust and that the thrust generation by the caudal fin may be enhanced by interception of the dorsal fin wake. Integration of experimental studies of fin wakes, computational approaches, and mechanical models of fin function promise understanding of instantaneous forces on fish fins during the propulsive cycle as well as exploration of a broader locomotor design space and its hydrodynamic consequences.     

The origins of adipose fins: an analysis of homoplasy and the serial homology of vertebrate appendages

Adipose fins are appendages found on the dorsal midline between the dorsal and caudal fins in more than 6000 living species of teleost fishes. It has been consistently argued that adipose fins evolved once and have been lost repeatedly across teleosts owing to limited function. Here, we demonstrate that adipose fins originated repeatedly by using phylogenetic and anatomical evidence. This suggests that adipose fins are adaptive, although their function remains undetermined. To test for generalities in the evolution of form in de novo vertebrate fins, we studied the skeletal anatomy of adipose fins across 620 species belonging to 186 genera and 55 families. Adipose fins have repeatedly evolved endoskeletal plates, anterior dermal spines and fin rays. The repeated evolution of fin rays in adipose fins suggests that these fins can evolve new tissue types and increased structural complexity by expressing fin-associated developmental modules in these new territories. Patterns of skeletal elaboration differ between the various occurrences of adipose fins and challenge prevailing hypotheses for vertebrate fin origin. Adipose fins represent a powerful and, thus far, barely studied model for exploring the evolution of vertebrate limbs and the roles of adaptation and generative biases in morphological evolution.     

Integration and modularity of teleostean pectoral fin shape and its role in the diversification of acanthomorph fishes

Phenotypic integration and modularity describe the strength and pattern of interdependencies between traits. Integration and modularity have been proposed to influence the trajectory of evolution, either acting as constraints or facilitators. Here, we examine trends in the integration and modularity of pectoral fin morphology in teleost fishes using geometric morphometrics. We compare the fin shapes of the highly diverse radiation of acanthomorph fishes to lower teleosts. Integration and modularity are measured using two‐block partial least squares analysis and the covariance ratio coefficient between the radial bones and lepidotrichia of the pectoral fins. We show that the fins of acanthomorph fishes are more tightly integrated but also more morphologically diverse and faster evolving compared to nonacanthomorph fishes. The main pattern of shape covariation in nonacanthomorphs is concordant with the main trajectory of evolution between nonacanthomorphs and acanthomorphs. Our findings support a facilitating role for integration during the acanthomorph diversification. Potential functional consequences and developmental mechanisms of fin integration are discussed.     

How predation shaped fish: the impact of fin spines on body form evolution across teleosts

It is well known that predators can induce morphological changes in some fish: individuals exposed to predation cues increase body depth and the length of spines. We hypothesize that these structures may evolve synergistically, as together, these traits will further enlarge the body dimensions of the fish that gape-limited predators must overcome. We therefore expect that the orientation of the spines will predict which body dimension increases in the presence of predators. Using phylogenetic comparative methods, we tested this prediction on the macroevolutionary scale across 347 teleost families, which display considerable variation in fin spines, body depth and width. Consistent with our predictions, we demonstrate that fin spines on the vertical plane (dorsal and anal fins) are associated with a deeper-bodied optimum. Lineages with spines on the horizontal plane (pectoral fins) are associated with a wider-bodied optimum. Optimal body dimensions across lineages without spines paralleling the body dimension match the allometric expectation. Additionally, lineages with longer spines have deeper and wider body dimensions. This evolutionary relationship between fin spines and body dimensions across teleosts reveals functional synergy between these two traits and a potential macroevolutionary signature of predation on the evolutionary dynamics of body shape.     

Ceratobregma striata,a new triplefin (Tripterygiidae) from northern Australia,and a record ofNorfolkia brachylepis from western Australia

A new species of tripterygiid fish, belonging to the recently described genusCeratobregma Holleman, 1987, is characterized by a total of 17 spines in the second dorsal fin, 8 rays in the third dorsal fin, 20 anal fin rays (spines+rays = 22), 14 + 21 lateral line scales, a total of 35–37 lateral scale series, small spiny scales on the sides of the head behind the eye and on the upper cheeks, 14 vertical dark stripes along the upper sides of the body, and 8 basal blotches on the second dorsal fin. A key to the species ofCeratobregma is presented.Norfolkia brachylepis is described and recorded from northwestern Australia.Norfolkia springen is synonymized withN. brachylepis.     

An early fossil remora (Echeneoidea) reveals the evolutionary assembly of the adhesion disc

The adhesion disc of living remoras (Echeneoidea: Echeneidae) represents one of the most remarkable structural innovations within fishes. Although homology between the spinous dorsal fin of generalized acanthomorph fishes and the remora adhesion disc is widely accepted, the sequence of evolutionary—rather than developmental—transformations leading from one to the other has remained unclear. Here, we show that the early remora †Opisthomyzon (Echeneoidea: †Opisthomyzonidae), from the early Oligocene (Rupelian) of Switzerland, is a stem-group echeneid and provides unique insights into the evolutionary assembly of the unusual body plan characteristic of all living remoras. The adhesion disc of †Opisthomyzon retains ancestral features found in the spiny dorsal fins of remora outgroups, and corroborates developmental interpretations of the homology of individual skeletal components of the disc. †Opisthomyzon indicates that the adhesion disc originated in a postcranial position, and that other specializations (including the origin of pectination, subdivision of median fin spines into paired lamellae, increase in segment count and migration to a supracranial position) took place later in the evolutionary history of remoras. This phylogenetic sequence of transformation finds some parallels in the order of ontogenetic changes to the disc documented for living remoras.