Comparative punting kinematics and pelvic fin musculature of benthic batoids

Although the majority of batoid elasmobranchs, skates and rays, are benthically associated, benthic locomotion has been largely overlooked in this group. Only skates have been previously described to perform a form of benthic locomotion termed “punting.” While keeping the rest of the body motionless, the skate's pelvic fins are planted into the substrate and then retracted caudally, which thrusts the body forward. In this study, we demonstrate that this form of locomotion is not confined to the skates, but is found across a range of phylogenetically and morphologically diverse batoid species. However, only the clearnose skate, Raja eglanteria, and the lesser electric ray, Narcine brasiliensis, performed “true punting,” in which only the pelvic fins were engaged. The yellow stingray, Urobatis jamaicensis, and the Atlantic stingray, Dasyatis sabina, performed “augmented punting,” in which pectoral fin movement was also used to generate thrust. Despite this supplemental use of pectoral fins, the augmented punters failed to exceed the punting capabilities of the true punters. The urobatid and the true punters all punted approximately half their disc length per punt, whereas the dasyatid punted a significantly shorter distance. The skate punted significantly faster than the other species. Examination of the pelvic fin musculature revealed more specialized muscles in the true punters than in the augmented punters. This concordance of musculature with punting ability provides predictive power regarding the punting kinematics of other elasmobranchs based upon gross muscular examinations. In contrast to previous assumptions, our results suggest that benthic locomotion is widespread among batoids. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Functional morphology of the pectoral fins in bamboo sharks, Chiloscyllium plagiosum: benthic vs. pelagic station-holding

Bamboo sharks (Chiloscyllium plagiosum) are primarily benthic and use their relatively flexible pectoral and pelvic fins to rest on and move about the substrate. We examined the morphology of the pectoral fins and investigated their locomotory function to determine if pectoral fin function during both benthic station-holding and pelagic swimming differs from fin function described previously in leopard sharks, Triakis semifasciata. We used three-dimensional kinematics and digital particle image velocimetry (DPIV) to quantify pectoral fin function in five white-spotted bamboo sharks, C. plagiosum, during four behaviors: holding station on the substrate, steady horizontal swimming, and rising and sinking during swimming. During benthic station-holding in current flow, bamboo sharks decrease body angle and adjust pectoral fin angle to shed a clockwise fluid vortex. This vortex generates negative lift more than eight times that produced during open water vertical maneuvering and also results in an upstream flow that pushes against the posterior surface of the pectoral fin to oppose drag. In contrast, there is no evidence of significant lift force in the wake of the pectoral fin during steady horizontal swimming. The pectoral fin is held concave downward and at a negative dihedral angle during steady horizontal swimming, promoting maneuverability rather than stability, although this negative dihedral angle is much less than that observed previously in sturgeon and leopard sharks. During sinking, the pectoral fins are held concave upward and shed a clockwise vortex with a negative lift force, while in rising the pectoral fin is held concave downward and sheds a counterclockwise vortex with a positive lift force. Bamboo sharks appear to sacrifice maneuverability for stability when locomoting in the water column and use their relatively flexible fins to generate strong negative lift forces when holding position on the substrate and to enhance stability when swimming in the water column.     

A Three-Dimensional Kinematics Analysis of a Koi Carp Pectoral Fin by Digital Image Processing

Pectoral fins fascinate researchers for their important role in fish maneuvers. By possessing a complicated flexible structure with several fin rays made by a thin film, the fin exhibits a three-dimensional (3D) motion. The complex 3D fin kinematics makes it challenging to study the performance of pectoral fin. Nevertheless, a detailed study on the 3D motion pattern of pectoral fins is necessary to the design and control of a bio-inspired fin rays. Therefore, a highspeed photography system is introduced in this paper to study the 3D motion of a Koi Carp by analyzing the two views of its pectoral fin simultaneously. The key motions of the pectoral fins are first captured in both hovering and retreating. Next, the 3D configuration of the pectoral fins is reconstructed by digital image processing, in which the movement of fin rays during fish retreating and hovering is obtained. Furthermore, the method of Singular Value Decomposition (SVD) is adopted to extract the basic motion patterns of pectoral fins from extensive image sequences, i.e. expansion, bending, cupping, and undulation. It is believed that the movement of the fin rays and the basic patterns of the pectoral fins obtained in the present work can provide a good foundation for the development and control of bionic flexible pectoral fins for underwater propeller.     

Functional Morphology of Aquatic Flight in Fishes: Kinematics, Electromyography, and Mechanical Modeling of Labriform Locomotion

Labriform locomotion is the primary swimming mode for many fishesthat use the pectoral fins to generate thrust across a broadrange of speeds. A review of the literature on hydrodynamics,kinematics, and morphology of pectoral fin mechanisms in fishesreveals that we lack several kinds of morphological and kinematicdata that are critical for understanding thrust generation inthis mode, particularly at higher velocities. Several needsinclude detailed three-dimensional kinematic data on speciesthat are pectoral fin swimmers across a broad range of speeds,data on the motor patterns of pectoral fin muscles, and thedevelopment of a mechanical model of pectoral fin functionalmorphology. New data are presented here on pectoral fin locomotionin Gomphosus varius, a labrid fish that uses the pectoral finsat speeds of 1 –6 total body lengths per second. Three-dimensionalkinematic data for the pectoral fins of G. varius show thata typical "drag-based" mechanism is not used in this species.Instead, the thrust mechanics of this fish are dominated bylift forces and acceleration reaction forces. The fin is twistedlike a propeller during the fin stroke, so that angles of attackare variable along the fin length. Electromyographic data onsix fin muscles indicate the sequence of muscle activity thatproduces antagonistic fin abduction and adduction and controlsthe leading edge of the fin. EMG activity in abductors and adductorsis synchronous with the start of abduction and adduction, respectively,so that muscle mechanics actuate the fin with positive work.A mechanical model of the pectoral fin is proposed in whichfin morphometrics and computer simulations allow predictionsof fin kinematics in three dimensions. The transmission of forceand motion to the leading edge of the fin depends on the mechanicaladvantage of fin ray levers. An integrative program of researchis suggested that will synthesize data on morphology, physiology,kinematics, and hydrodynamics to understand the mechanics ofpectoral fin swimming.     

Ecomorphology of Locomotion in Labrid Fishes

The Labridae is an ecologically diverse group of mostly reef associated marine fishes that swim primarily by oscillating their pectoral fins. To generate locomotor thrust, labrids employ the paired pectoral fins in motions that range from a fore-aft rowing stroke to a dorso-ventral flapping stroke. Species that emphasize one or the other behavior are expected to benefit from alternative fin shapes that maximize performance of their primary swimming behavior. We document the diversity of pectoral fin shape in 143 species of labrids from the Great Barrier Reef and the Caribbean. Pectoral fin aspect ratio ranged among species from 1.12 to 4.48 and showed a distribution with two peaks at about 2.0 and 3.0. Higher aspect ratio fins typically had a relatively long leading edge and were narrower distally. Body mass only explained 3% of the variation in fin aspect ratio in spite of four orders of magnitude range and an expectation that the advantages of high aspect ratio fins and flapping motion are greatest at large body sizes. Aspect ratio was correlated with the angle of attachment of the fin on the body (r = 0.65), indicating that the orientation of the pectoral girdle is rotated in high aspect ratio species to enable them to move their fin in a flapping motion. Field measures of routine swimming speed were made in 43 species from the Great Barrier Reef. Multiple regression revealed that fin aspect ratio explained 52% of the variation in size-corrected swimming speed, but the angle of attachment of the pectoral fin only explained an additional 2%. Labrid locomotor diversity appears to be related to a trade-off between efficiency of fast swimming and maneuverability in slow swimming species. Slow swimmers typically swim closer to the reef while fast swimmers dominate the water column and shallow, high-flow habitats. Planktivory was the most common trophic associate with high aspect ratio fins and fast swimming, apparently evolving six times.     

Allometric growth of the trunk leads to the rostral shift of the pelvic fin in teleost fishes

The pelvic fin position among teleost fishes has shifted rostrally during evolution, resulting in diversification of both behavior and habitat. We explored the developmental basis for the rostral shift in pelvic fin position in teleost fishes using zebrafish (abdominal pelvic fins) and Nile tilapia (thoracic pelvic fins). Cell fate mapping experiments revealed that changes in the distribution of lateral plate mesodermal cells accompany the trunk-tail protrusion. Presumptive pelvic fin cells are originally located at the body wall adjacent to the anterior limit of hoxc10a expression in the spinal cord, and their position shifts rostrally as the trunk grows. We then showed that the differences in pelvic fin position between zebrafish and Nile tilapia were not due to changes in expression or function of gdf11. We also found that hox-independent motoneurons located above the pelvic fins innervate into the pelvic musculature. Our results suggest that there is a common mechanism among teleosts and tetrapods that controls paired appendage positioning via gdf11, but in teleost fishes the position of prospective pelvic fin cells on the yolk surface shifts as the trunk grows. In addition, teleost motoneurons, which lack lateral motor columns, innervate the pelvic fins in a manner independent of the rostral-caudal patterns of hox expression in the spinal cord.     

Pelvic girdle shape predicts locomotion and phylogeny in batoids

In terrestrial vertebrates, the pelvic girdle can reliably predict locomotor mode. Because of the diminished gravitational effects on positively buoyant bony fish, the same relationship does not appear to exist. However, within the negatively buoyant elasmobranch fishes, benthic batoids employ pelvic fin bottom‐walking and punting as primary or supplementary forms of locomotion. Therefore, in this study, we employed geometric and linear morphometrics to investigate if their pelvic girdles exhibit shape characteristics similar to those of sprawling terrestrial vertebrates. We tested for correlates of pelvic girdle shape with 1) Order, 2) Family, 3) Swim Mode, and/or 4) Punt Mode. Landmarks and semilandmarks were placed along outlines of dorsal views of 61 batoid pelvic girdles (3/3 orders, 10/13 families, 35/72 genera). The first three relative warps explained 88.45% of the variation among individuals (P < 0.01%). Only Order and Punt Mode contained groups that were all significantly different from each other (P < 0.01%). Discriminant function analyses indicated that the majority of variation within each category was due to differences in extension of lateral and prepelvic processes and puboischiac bar angle. Over 60% of the original specimens and 55% of the cross‐validated specimens were correctly classified. The neutral angle of the propterygium, which articulates with the pelvic girdle, was significantly different among punt modes, whereas only pectoral fin oscillators had differently shaped pelvic girdles when compared with batoids that perform other swimming modes (P < 0.01). Pelvic girdles of batoids vary greatly, and therefore, likely function in ways not previously described in teleost fishes. This study illustrates that pelvic girdle shape is a good predictor of punt mode, some forms of swimming mode, and a species' Order. Such correlation between locomotor style and pelvic girdle shape provides evidence for the convergent evolution of morphological features that support both sprawled‐gait terrestrial walking and aquatic bottom‐walking. J. Morphol. 275:100–110, 2014. © 2013 Wiley Periodicals, Inc.     

Work that body: fin and body movements determine herbivore feeding performance within the natural reef environment

Herbivorous fishes form a keystone component of reef ecosystems, yet the functional mechanisms underlying their feeding performance are poorly understood. In water, gravity is counter-balanced by buoyancy, hence fish are recoiled backwards after every bite they take from the substrate. To overcome this recoil and maintain contact with the algae covered substrate, fish need to generate thrust while feeding. However, the locomotory performance of reef herbivores in the context of feeding has hitherto been ignored. We used a three-dimensional high-speed video system to track mouth and body kinematics during in situ feeding strikes of fishes in the genus Zebrasoma, while synchronously recording the forces exerted on the substrate. These herbivores committed stereotypic and coordinated body and fin movements when feeding off the substrate and these movements determined algal biomass removed. Specifically, the speed of rapidly backing away from the substrate was associated with the magnitude of the pull force and the biomass of algae removed from the substrate per feeding bout. Our new framework for measuring biting performance in situ demonstrates that coordinated movements of the body and fins play a crucial role in herbivore foraging performance and may explain major axes of body and fin shape diversification across reef herbivore guilds.     

Use of biorobotic models of highly deformable fins for studying the mechanics and control of fin forces in fishes

Bony fish swim with a level of agility that is unmatched in human-developed systems. This is due, in part, to the ability of the fish to carefully control hydrodynamic forces through the active modulation of the fins' kinematics and mechanical properties. To better understand how fish produce and control forces, biorobotic models of the bluegill sunfish's (Lepomis macrochirus) caudal fin and pectoral fins were developed. The designs of these systems were based on detailed analyses of the anatomy, kinematics, and hydrodynamics of the biological fins. The fin models have been used to investigate how fin kinematics and the mechanical properties of the fin-rays influence propulsive forces and to explore kinematic patterns that were inspired by biological motions but that were not explicitly performed by the fish. Results from studies conducted with the fin models indicate that subtle changes to the kinematics and mechanical properties of fin rays can significantly impact the magnitude, direction, and time course of the 3D forces used for propulsion and maneuvers. The magnitude of the force tends to scale with the fin's stiffness, but the direction of the force is not invariant, and this causes disproportional changes in the magnitude of the thrust, lift, and lateral components of force. Results from these studies shed light on the multiple strategies that are available to the fish to modulate fin forces.     

Ontogenetic change in novel functions: waterfall climbing in adult Hawaiian gobiid fishes

Juveniles from three species of Hawaiian gobiid fishes climb waterfalls as part of an amphidromous life cycle, allowing them to re-penetrate adult upstream habitats after being swept out to the ocean upon hatching. The importance of climbing for juvenile stream gobies is well established, but adult fish in upstream island habitats also face potential downstream displacement by periodic disturbances. Thus, retention of climbing ability could be advantageous for adult stream gobies. Climbing performance might be expected to decline among adults, however, due to the tendency for mass-specific muscular power production to decrease with body size, and a lack of positively allometric growth among structures like the pelvic sucker that support body weight against gravity. To evaluate changes in waterfall-climbing ability with body size in Hawaiian stream gobies, we compared climbing performance and kinematics between adults and juveniles from three species: Awaous guamensis , Sicyopterus stimpsoni and Lentipes concolor . For species in which juveniles climbed using 'powerbursts' of axial undulation, adult performance and kinematics showed marked changes: adult A. guamensis failed to climb, and adult L. concolor used multiple pectoral fin adductions to crutch up surfaces at slow speeds, rather than rapid powerbursts. Adult S. stimpsoni , like juveniles, still used oral and pelvic suckers to 'inch' up surfaces and climbed at speeds comparable to those of juveniles. However, unlike juveniles, adult S. stimpsoni also add pectoral fin crutching to every climbing cycle. Thus, although powerburst species appear to be particularly susceptible to size-related declines in waterfall-climbing performance, the addition of compensatory mechanisms prevents the loss of this novel function in some species.     

Development of the paired fins in the paddlefish, Polyodon spathula

In Polyodon spathula, the pectoral fin radials, with the exception of the metapterygium, are derived from the decomposition of a single continuous cartilage fin plate that is continuous with the scapulocoracoid. This cartilage sheet develops two interior splits to form three precursor pieces, and these decompose in a predictable way to generate the propterygium and radials. The metapterygium is an extension of the scapulocoracoid that segments off of it during early development. To our knowledge, this has not been reported for acipenserids or other basal actinopterygians. In teleosts, the proximal radials also develop from the "break up" of an initially continuous paddle-like sheet of cartilage along the posterior edge of the scapulocoracoid, and in Polypterus and sharks a similar pattern holds. Thus, the pattern observed in Polyodon may represent the basal developmental condition for the gnathostome pectoral fin. The process underlying development of the superficially similar cartilages of the pelvic and pectoral fins is different. In the pectoral fin, the metapterygium is segmented off of the scapulocoracoid and other radials form from the decomposition of the cartilage plate. In contrast, individual rod-like basipterygial elements form in a close one-to-one correspondence with the middle radials of the pelvic fin, but later fuse to form an anterior element that is branched in appearance. To evaluate further claims of similarity among the pectoral and pelvic fin elements of various fishes, the course of the development of these structures must be observed. The pectoral fin and girdle in Polyodon ossifies in a different sequence than that proposed as ancestral (and highly conserved) for actinopterygians: the supracleithrum ossifies significantly before the cleithrum. The later ossification of the cleithrum in Polyodon may be related to the primary use of the caudal fin vs. the pectoral fins in their locomotion.     

A new fish species Hemibrycon (Characiformes: Characidae)

Hemibrycon pautensis (Characiformes, Characidae), a new fish species from Paute River, eastern Ecuador is described. Diagnostic characteristics: eight to nine branched rays in the dorsal fin (vs. six to seven), and 27 - 28 in the anal fin (vs. 16 - 26, except in H. dariensis which presents 22 - 27, in H. metae 26 - 31 and H. jabonero 23 - 28); a no occurrence of dorsal pharyngeal plate (vs. occurrence); a cartilaginous and divided-in-two basihial (vs. an osseous base and a cartilaginous upper part). Hemibrycon pautensis resembles H. metae by its oblique external edge of the pelvic fins. They can be distinguished by the position of the pectoral fins in relation to the snout (38.24-41.6% in H. pautensis vs. 21.21-25.87) and by the position of the pectoral fins in relation to the origin of the dorsal fin (20.95-24.30 in Hemibrycon pautensis vs. 35.89-42.63), and by the number of proximate radials in the pectoral girdle (five in Hemibrycon pautensis vs. three to four). In addition, the geographic distribution of H. metae is restricted to the upper part of the Meta River in Colombia and can be distinguished of H. boquiae by: the number of scales between the lateral-line and the origin of the dorsal fin (eight in H. pautensis vs. 5-7); the distance between the snout and the pelvic fins (38.00-42.90 % in H. pautensis vs. 42.9-46.19%); the pelvic fins length (13.77-17.96% in H. pautensis vs. 10.72-13.21%); and the snout length (21.34-27.88 in H. pautensis vs. 26.92-33.66%).     

Internal morphology and function of paired fins in the epaulette shark,hemiscyllium ocellatum

Paired fins and associated internal structures of the epauletic sharkHemiscyllium ocellatum, were described on the basis of three specimens. A comparison with other genera showed the epaulette shark to be, characterized by two elongated basal cartilages articulating with a distally projecting articular condyle on the coracoid, a loosely separated radial series with an intermediate series, a levator pectoralis inferior muscle and an anterolaterally developed depressor pectoralis muscle in the pectoral fin, and an elongated anterior pelvic basal cartilage articulating with a distally projecting articular condyle and an anterolaterally developed depressor pelvicus muscle in the pelvic fin. In captivity, the sharks exhibited both upright and crawling behavior on the bottom by using the pectoral and pelvic fins and bending the body. The distinctive morphological characters are shared by otherHemiscyllium species and are suggested as important factors enabling their unique behavior associated with a complex coral reef habitat.     

Larval development of Argentine hake Merluccius hubbsi

The ontogeny of the developmental stages of the hake Merluccius hubbsi is described. Fish larvae and post-transitional juveniles were collected in the Nor-Patagonian area from 1989 to 2004. The opening of the mouth and the pigmentation of the eyes are coincident with yolk resorption, finishing the yolk-sac stage. This species presents pigmentation on the head, trunk and tail typical of gadiform larvae. Pectoral fin development is completed during the transformation stage. The post-transitional juvenile stage begins when the fin-ray complements are complete and squamation begins. The fins become fully formed in the following sequence: pelvic fins, first dorsal fin, second dorsal and anal fins together, caudal fin and pectoral fins. The caudal complex is totally developed in larvae of 22·0–23·0 mm standard lengths ( L _S) and all vertebral elements are first observed in larvae of 8·5 mm L _S. The rate of development of M. hubbsi observed in this study could be faster than the rates reported for other species of Merluccius by different authors.     

Biomechanics of swimming in the pufferfish Diodon holocanthus: propulsive momentum enhancement is an adaptation for thrust production in an undulatory median and paired-fin swimmer

A form of large-amplitude elongated-body theory appropriate for the analysis of undulatory fins attached to a rigid body of elliptical section suggests a benefit due to momentum enhancement relative to the fins on their own. This theoretical prediction is experimentally confirmed for the first time. Theoretical momentum enhancement factors for Diodon holocanthus (2.2 and 2.7 for the median and pectoral fins, respectively) compared well to inferred thrust values determined from particle-image velocimetry (PIV) wake measurements (2.2-2.4 and 2.7-2.9). Caudal fin mean theoretical thrust was not significantly different from measured (PIV) values (n = 24, P > 0.05), implying no momentum enhancement. Pectoral-fin thrust was half that of the median and caudal fins due to high fin-jet angles, low circulation and momentum. Average total fin thrust and fish drag were not significantly different (n = 24, P > 0.05). Vortex rings generated by the fins were elliptical, with size dependent on fin chord and stroke amplitude. Hydrodynamic advantages (thrust enhancement at no cost to hydrodynamic efficiency, reduction of side forces minimizing energy wasting yawing motions and body drag) are probably common among rigid-bodied organisms propelled by undulatory fins. A trade-off between momentum enhancement and the rate of momentum generation (thrust force) sets a practical limit to the former. For small fins whilst momentum enhancement is high, absolute thrust is low. In addition, previously suggested limitations on thrust enhancement set by reductions in propulsive force associated with progressive reductions in fin wavelength are found to be biologically unrealistic.     

Life in the flow lane: differences in pectoral fin morphology suggest transitions in station-holding demand across species of marine sculpin

Aquatic organisms exposed to high flow regimes typically exhibit adaptations to decrease overall drag and increase friction with the substrate. However, these adaptations have not yet been examined on a structural level. Sculpins (Scorpaeniformes: Cottoidea) have regionalized pectoral fins that are modified for increasing friction with the substrate, and morphological specialization varies across species. We examined body and pectoral fin morphology of 9 species to determine patterns of body and pectoral fin specialization. Intact specimens and pectoral fins were measured, and multivariate techniques determined the differences among species. Cluster analysis identified 4 groups that likely represent differences in station-holding demand, and this was supported by a discriminant function analysis. Primarily, the high-demand group had increased peduncle depth (specialization for acceleration) and larger pectoral fins with less webbed ventral rays (specialization for mechanical gripping) compared to other groups; secondarily, the high-demand group had a greater aspect ratio and a reduced number of pectoral fin rays (specialization for lift generation) than other groups. The function of sculpin pectoral fins likely shifts from primarily gripping where demand is likely low, to an equal dependence on gripping and negative lift generation where demand is likely high. Specialization of the ventral pectoral fin region for gripping likely contributes to the recent diversification of some species into high-demand habitats.     

Larval development of Hypsophrys nicaraguensis (Pisces: Cichlidae) under laboratory conditions

The cichlid Hypsophrys nicaraguensis is a popular fish known as butterfly, and despite its widespread use as pets, little is known about its reproductive biology. In order to contribute to this knowledge, the study describes the relevant larval development characteristics, from adult and larval cultures in captivity. Every 12h, samples of larvae were collected and observed under the microscope for larval stage development, and every 24h morphometric measurements were taken. Observations showed that at 120h, some larvae had swimming activity and the pectoral fins development was visible; at 144h, the dorsal fin appear and all larvae started food intake; at 168h, the formation of anal fins begins, small rudiments of pelvic fins emerge, the separation of caudal fin from anal and dorsal fins starts, and the yolk sac is reabsorbed almost completely; at 288h, the pelvic fins starts to form; at 432h, the rays and spines of dorsal and anal fins can be distinguished, both the anal and the dorsal fins have the same number of spines and rays as in adults. After 480h larvae have the first scales, ending the larval stages and starting the transformation to fingerlings. Larvae were successfully fed with commercial diet.     

Acanthodes bourbonensis n.sp., ein neuer Acanthodier (Acanthodii: Pisces) aus dem Autunium (Unterperm) von Bourbon l’Archambault (Allier,Frankreich)

WithAcanthodes bourbonensis n.sp. another acanthodian from Lower Permian basins of Europe is described. The new species is similar toAcanthodes gracilis (Beyrich) from Silesia (Poland), but it differs from this and all other species of the genus in the development of the pectoral fins, dorsal fin, anal fin and caudal fin. In pectoral fins, dorsal and anal fin there are different ceratotrichia as supporting elements and pectoral fins are attaching along a row of oblonged large scales. In the caudal fin there is an epichoral appendix first found byHeyler (1969).