Lunate Tails Research Articles

The fast and the tendinous? Locomotor modifications of the caudal peduncle in Gila spp. from the American Southwest

In the American Southwest, the fishes within the genus Gila evolved in an environment with seasonal rainstorms that caused stochastic flooding. Some species within this genus, such as bonytail (Gila elegans), possess locomotor morphologies that are similar to those seen in high-performance swimmers such as tuna and lamnid sharks. These shared features include a shallow caudal peduncle, lunate tail, and mechanisms to transmit force from the anterior musculature to the tail fin. We compared the skeletal anatomy of the caudal region of bonytail to roundtail chub (Gila robusta) and humpback chub (Gila cypha) to determine which vertebral elements have been modified to create a shallow peduncle. We also tested the tensile strength of the red (slow oxidative) axial muscle by performing a standard stress test. If the muscle can withstand a large load, this suggests it may play a tendon-like role in transmitting force from the anterior muscle to the hypural plate of the tail. Lastly, we measured the collagen content of the red axial muscle (visualized using serial sections and Masson’s trichrome stain) to determine if increased tensile strength is associated with increased collagen content. We found bonytail caudal peduncles are characterized by acute vertebral spines and have red axial muscle that can resist tearing under tension. Roundtail chub peduncles are characterized by relatively more obtuse angles and the red muscle tears easily under tension. Humpback chub possess an intermediate morphology, with relatively obtuse vertebral spine angles and the red muscle can resist tearing under tension. Bonytail have increased collagen content in posterior red axial muscle compared to the anterior musculature also suggesting a tendon-like role of the posterior red muscle. In combination with previous studies of swimming performance, our findings suggest that the axial musculature of bonytail may play a role in transmitting force directly to the shallow peduncle in a manner similar to that of the great lateral tendon of scombrids.

Flexibility of Heterocercal Tails: What Can the Functional Morphology of Shark Tails Tell Us about Ichthyosaur Swimming?

SynopsisThe similarities between ichthyosaurs and sharks are a text-book example of convergence, and similarities in tail morphology have led many to theorize that they had similar swimming styles. The variation of ichthyosaur tail shapes is encompassed within the diversity of shark families. In particular early ichthyosaurs have asymmetrical tails like the heterocercal tails of carcharhinid sharks, while later occurring ichthyosaurs have lunate tails similar to those of lamnid sharks. Because it is not possible to measure ichthyosaur tail function, the goal of this study is to measure and compare the flexibility and stiffness of lunate and heterocercal shark tails, and to measure skeletal and connective tissue features that may affect tail flexibility. We measured flexibility in 10 species and focused on five species in particular, for dissection: one pelagic and one bottom-associated individual from each order, plus the common thresher shark (Alopias vulpinus), a tail-slapping specialist. As expected, lunate tails were overall less flexible than heterocercal tails and had greater flexural stiffness. Our results suggest that the cross-sectional profile of the skeletally supported dorsal lobe dictates flexural stiffness, but that changing tissue composition dictates flexural stiffness in the ventral lobe. We also found structural differences that may enable the tail slapping behavior of the common thresher shark. Finally, we discuss how our morphological measurements compare to ichthyosaur measurements from the literature; noting that similarities in functional morphology suggest sharks may be a good analog for understanding ichthyosaur swimming biomechanics.

Topology Optimization of the Caudal Fin of the Three-Dimensional Self-Propelled Swimming Fish

AbstractBased on the boundary vorticity-flux theory, topology optimization of the caudal fin of the three-dimensional self-propelled swimming fish is investigated by combining unsteady computational fluid dynamics with moving boundary and topology optimization algorithms in this study. The objective functional of topology optimization is the function of swimming efficiency, swimming speed and motion direction control. The optimal caudal fin, whose topology is different from that of the natural fish caudal fin, make the 3D bionic fish achieve higher swimming efficiency, faster swimming speed and better maneuverability. The boundary vorticity-flux on the body surface of the 3D fish before and after optimization reveals the mechanism of high performance swimming of the topology optimization bionic fish. The comparative analysis between the swimming performance of the 3D topology optimization bionic fish and the 3D lunate tail bionic fish is also carried out, and the wake structures of two types of bionic fish show the physical nature that the swimming performance of the 3D topology optimization bionic fish is significantly better than the 3D lunate tail bionic fish.

Numerical studies on locomotion perfromance of fish-like tail fins

Flapping plates of typical fishlike tail shapes are simulated to investigate their locomotion performance using the multi-block Lattice Boltzmann Method (LBM) and Immersed Boundary (IB) method. Numerical results show that fishlike forked configurations have better locomotion performance compared with unforked plates. Based on our results, the caudal fin in carangi-form mode has greater thrust, and the lunate tail fin in thunniform mode has higher efficiency. These findings are qualitatively consistent with biological observations of fish swimming. Analysis of wake topology shows that the wake of the forked plate consists of a chain of alternating reverse horseshoe-like vortical structures. These structures induce a backward jet and generate a positive thrust. Moreover, this backward jet has a more favorable direction compared with that behind an unforked plate.

Fuzzy Control of Biomimetic Propulsion Using Flapping Foil

Flapping lunate tail is studied through hydrodynamic test in water channel,and proper vorticity control methods for flapping foil propulsion are discussed based on the principal vorticity parameters,such as Strouhal number(St),angle of attack,the dimensionless transverse amplitude,and phase difference.Free-swimming test is carried out on an unmanned underwater vehicle with flapping tail.The results show that power consumption of vehicle can be reduced effectively through active control of St and other vorticity parameters using fuzzy control methods,and greater swimming speed can be obtained.The maximum swimming speed as high as 1.87 m/s is measured.Meanwhile,St of flapping tail is controlled within the range of 0.4～0.5.

Fuzzy Vorticity Control of a Biomimetic Robotic Fish Using a Flapping Lunate Tail

Vorticity control mechanisms for flapping foils play a guiding role in both biomimetic thrust research and modeling the forward locomotion of animals with wings, fins, or tails. In this paper, a thrust-producing flapping lunate tail is studied through force and power measurements in a water channel. Proper vorticity control methods for flapping tails are discussed based on the vorticity control parameters: the dimensionless transverse amplitude, Strouhal number, angle of attack, and phase angle. Field tests are conducted on a free-swimming biomimetic robotic fish that uses a flapping tail. The results show that active control of Strouhal number using fuzzy logic control methods can efficiently reduce power consumption of the robotic fish and high swimming speeds can be obtained. A maximum speed of 1.17 length specific speed is obtained experimentally under conditions of optimal vorticity control. The St of the flapping tail is controlled within the range of 0.4∼0.5.

Two new cichild species Neolamprologus (Teleostei: Cichlidae) from Lake Tanganyika, East Africa

Two new cichlid species, Neolamprologus walteri sp. nov. and N. chitamwebwai sp. nov., from the Bangwe peninsula (Tanzania), on the east coast of Lake Tanganyika, are described. Both species belong to the N. savoryi species complex, characterised by a lunate tail with long filaments. Neolamprologus chitamwebwai and N. walteri are highly stenotopic species, restricted to rocky habitat. They occur sympatrically with each other and with two other species of the complex but occupy different microhabitats. Neolamprologus walteri lives in large numbers in sheltered areas with rubble substrate whereas N. chitamwebwai occurs in much lower numbers on more exposed parts of the rocky shores, with large boulders, lower sedimentation rates, coarser sediment, and higher visibility. The two new species although closely related, show clear ecological differentiation. Evidence of ecological differentiation between closely related sympatric cichlids is rare in literature. Closely related sympatric cichlids in the large African lakes (with literature mainly referring to Lake Victoria and Lake Malawi) have been suggested to differ little in diet and habitat use, which has encouraged the idea that cichlid species can coexist without niche partitioning. Our paper provides a different perspective with cichlids from Lake Tanganyika.

Speed limits on swimming of fishes and cetaceans

Physical limits on swimming speed of lunate tail propelled aquatic animals are proposed. A hydrodynamic analysis, applying experimental data wherever possible, is used to show that small swimmers (roughly less than a metre long) are limited by the available power, while larger swimmers at a few metres below the water surface are limited by cavitation. Depending on the caudal fin cross-section, 10-15 m s(-1) is shown to be the maximum cavitation-free velocity for all swimmers at a shallow depth.

Adaptive Control of Shape Memory Alloy Actuators for Underwater Biomimetic Applications

In actuator technology active or smart materials have opened up new horizons in terms of actuation simplicity, compactness, and miniaturization potential. One such material is the nickel-titanium shape memory alloy (NiTi SMA), which is gaining widespread use in a variety of applications. The numerous advantages of SMA over traditional actuators are of particular interest in the area of underwater vehicle design, particularly the development of highly maneuverable vehicles of a design based on the swimming techniques and anatomic structure of e sh. An SMA actuation cycle consists of heating/cooling half-cycles, currently imposing a limit on the frequency of actuation to well below 1 Hz in air because of slow cooling. The aquatic environment of underwater vehicles lends itself to cooling schemes that use the excellent heat-transfer properties of water, thus enabling much higher actuation frequencies. A controller for SMA actuators must account not only for large hysteretic nonlinearities betweenactuatoroutput (strainordisplacement )andinput(temperature ),butalso thethermalcontrolforresistive heating via an applied current. The control of SMA in water presents a problem not encountered when actuating in air: accurate temperature feedback for the SMA is very dife cult in water. We overcome this problem by using a simplie ed thermal model to estimate the temperatureof the wire in conjunction with an adaptivehysteresismodel, which relates the actuator output to the estimated temperature. Experimental results are provided, showing that this method for control of an SMA wire works equally well both in air and in water, with only rough estimates (easily obtained )ofthethermal parameters.Successful tracking of referencedisplacementsignals with frequencies up to 2 Hz and relatively large amplitudes have been demonstrated experimentally. I. Introduction I N aerodynamicsand hydrodynamics birds and e sh have inspired and guided the development of aircraft and underwater vehicles. These manmademachinesseemsoprimitive compared to their natural counterparts in terms of intelligence, efe ciency, agility, adaptability, and functionalcomplexity. These and other similar observationsandissuesthathavebeenaddressedbythescientie ccommunity havetriggered theformulation of thescience ofbiomimetics and have inspired new approaches to old problems. In the area of underwater vehicle design, the development of highly maneuverable vehicles is presently of interest, with their design being based on the swimming techniques and anatomic structure of e sh; primarily the undulatory body motions, the highly controllable e ns, and the large aspect ratio lunate tail. The tailoring and implementation of the accumulated knowledge into biomimetic vehicles is a task of multidisciplinary nature with two of the dominant e elds being

Hydrodynamic characteristics of a lunate shape oscillating propulsor

Research was conducted to study the hydrodynamic efficiency of a foil with aft-swept wing tips. A potential flow based time domain panel method was formulated to predict the performance of a lunate and rectangular foil in large amplitude, unsteady motion. Skin drag was approximated and boundary layer growth and separation were also estimated. Hydrodynamic efficiency was evaluated in terms of propulsive efficiency and thrust coefficient of the foil. Results are presented for a lunate shaped planform and for a rectangular foil. Predictions show that the lunate shaped planform has a substantially higher propulsive efficiency (13% higher) than the rectangular foil under heavy load conditions when the feathering parameter is zero, throughout a range of reduced frequencies (0.2 to 1.8). Under a medium load condition, however, the rectangular foil gave a higher propulsive efficiency at reduced frequencies less than 0.5 and the same efficiency value at a reduced frequency of 1.8. For a practical range of reduced frequencies between 0.5 and 1.0, the lunate tail gave higher propulsive efficiency. The lunate planform gave a lower thrust coefficient at a heavy load and higher thrust at a medium load condition than the rectangular planform for all reduced frequencies.

Eel-like swimming in the earliest ichthyosaurs

ICHTHYOSAURS are extinct marine reptiles, probably belonging to the Diapsida1, that ranged from the Early Triassic to Late Cretaceous2,3. Post-Triassic ichthyosaurs achieved the highest level of aquatic adaptation among reptiles4, with a streamlined body, lunate tail and a dorsal fin, features exemplified today by thunniform (tuna-like) fishes. However, little is known of how such a body plan evolved from a terrestrial diapsid. Here we report the most complete specimen of the oldest known ichthyosaur, Chensaurus, representing a transition between the two body plans. The specimen, which has a partial skin impression, has a small caudal fin, a long and narrow body, and a high presacral vertebral count. These features all suggest an anguilliform swimming mode. Later ichthyosaurs retained the high vertebral count, but overcame the high swimming costs of this plesiomorphy, achieving a rigid tunniform bauplan by evolving discoidal vertebrae, and a deep fusiform body. Chensaurus therefore seems to be an evolutionary intermediate between the shorter-bodied terrestrial stock from which the group evolved, and advanced thunniform ichthyosaurs.

The affinities and ecology of Triassic ichthyosaurs

The resolution of the problem of ichthyosaurian affinities has been complicated and prolonged by an emphasis on the morphology of Jurassic ichthyosaurs, some 30 to 40 m.y. younger than the earliest known species. Morphology of the skull and axial skeleton of Triassic ichthyosaurs is the key to resolving the problem. The ichthyosaurs are included here in the Subclass Diapsida on the basis of skull roof and palatal features. Within the Diapsida, ichthyosaurs appear to be most closely related to the Younginiformes of the Infraclass Lepidosauromorpha, with which they may have shared a common ancestor. Triassic ichthyosaurs were considerably different ecologically from their Jurassic descendants. Many Triassic ichthyosaurs were longer bodied and had long, broad tails in contrast to the deep, streamlined bodies and lunate tails of Jurassic species. As a result, Triassic ichthyosaurs probably tended to be ambush predators, whereas Jurassic ichthyosaurs were mainly pursuit predators. Early ichthyosaurs may have been more generalized in their prey preference, as suggested by heterodonty in many Early and Middle Triassic species.

Lunate-tail swimming propulsion. Part 2. Performance analysis

The theory of an oscillating, high-aspect-ratio, lifting surface with a curved centreline (Cheng & Murillo 1984) is applied to a performance analysis of lunate-tail swimming propulsion. Thrust, power and propulsive efficiency are calculated for model lunate tails with various combinations of mode shapes and morphological features to ascertain the viability of the proportional-feathering concept, and to determine the influence of sweep and centreline curvature. One of the principal conclusions concerns the interchangeability of the heaving amplitude of the peduncle (identified with the major pitching axis) with the centreline sweep, and its effect on the propulsive efficiency, while maintaining the same thrust. Hydrodynamic reasons are also offered for the apparent preference for the crescent-moon fin shape over the V-shape at moderate sweep angles, and for the large sweep angles often found in V-shaped fins.

Optimum pitching axes in flapping wing propulsion

The locomotion of many birds, insects, marine mammals and fish with lunate tails is characterized by a heaving and pitching motion of a high aspect ratio wing or fin. The time dependent fluid forces, moment of force and rate of work done by the propulsive appendages can be estimated using two-dimensional unsteady aerodynamic theories. For a prescribed motion, the moment and power are further analyzed to find the axes for which the mean square moment is minimum, the mean power to maintain the moment is zero, the mean square power to maintain moment is minimum, and the mean square power to maintain lift equals the mean square power. In this way, several new axes of physiological significance are identified, and their positions are found to depend on the beat frequency and the wing loading. This choice of pitching axis may enhance performance and control of rapid maneuvers and thus enable the organism to more adeptly cope with turbulent environmental conditions, to avoid danger or to more easily capture food. Several observations and measurements on wings in nature support the results of this analysis.

Maneuverability of the Humboldt penguin (Spheniscus humboldti) during swimming

Analyses of high-speed film of captive Humboldt penguins (Spheniscus humboldti) show that there is no correlation between the radii of submerged turns and swimming speed. The sharpest turns had a mean turn radius (n = 5) of 0.14 m (0.24 body lengths), were powered by beating wings, and used multiple steering structures (beak, tail, feet, wings) and a flexing body. This turn radius is similar to that of fast-swimming fish with rigid bodies and lunate tails but is greater than that of slower fish with flexible bodies. The maximum turning rate was 10.05 radians s−1. Maneuverability is augmented by neck-extending strike behavior and porpoising leaps.

Lunate-tail swimming propulsion as a problem of curved lifting line in unsteady flow. Part 1. Asymptotic theory

The asymptotic theory of a high-aspect-ratio wing in an incompressible flow is generalized to an oscillating lifting surface with a curved centreline in the domain where the reduced frequency based on the half-span is of order unity. The formulation allows applications to lightly loaded models of lunate-tail swimming and ornithopter flight, provided that the heaving displacement does not far exceed the mean wing chord. The analysis include the quasi-steady limit, in which the crescent-moon wing problem considered earlier by Thurber (1965) is solved and several aerodynamic properties of swept wings are explained.Among the important three-dimensional and unsteady effects are corrections for the centreline curvature and for the spanwise components of the locally shed vortices. Comparison of the lift distributions obtained for model lunate tails with data computed from the doublet-lattice method (Albano & Rodden 1969) lends support to the asymptotic theory.

The unsteady quasi-vortex-lattice method with applications to animal propulsion

In the early theoretical study of aquatic animal propulsion either the two-dimensional theory or the large aspect-ratio theory has been generally used. Only recently has the unsteady lifting-surface theory with the continuous loading approach been applied to the study of this problem by Chopra & Kambe (1977). Since it is well known that the continuous loading approach is difficult to extend to general configurations, a new quasi-continuous loading method, applicable to general configurations and yet accurate enough for practical applications, is developed in this paper. The method is an extension of the steady version of Lan (1974) and is particularly suitable for predicting the unsteady lead-edge suction during harmonic motion.The method is applied to the calculation of the propulsive efficiency and thrust for some swept and rectangular planforms by varying the phase angles between the pitching and heaving motions. It is found that with the pitching axis passing through the trailing edge of the root chord and the reduced frequency k equal to 0·75 the rectangular planform is quite sensitive in performance to the phase angles and may produce drag instead of thrust. These characteristics are not shared by the swept planforms simulating the lunate tails. In addition, when the pitching leads the heaving motion by 90°, the phase angle for nearly maximum efficiency, the planform inclination caused by pitching contributes to the propulsive thrust over a large portion of the swept planform, while, for the rectangular planform, only drag is produced from the planform normal force at k = 0·75. It is also found that the maximum thrust is not produced with maximum efficiency for all planforms considered. The theory is then applied to the study of dragonfly aerodynamics. It is shown that the aerodynamically interacting tandem wings of the dragonfly can produce high thrust with high efficiency if the pitching is in advance of the flapping and the hindwing leads the forewing with some optimum phase angle. The responsible mechanism allows the hindwing to extract wake energy from the forewing.

Hydromechanics of lunate-tail swimming propulsion. Part 2

This paper investigates the propulsive performance of the lunate tails of aquatic animals achieving high propulsive efficiency (the hydromechanical efficiency being defined as the ratio of the work done by the mean forward thrust to the mean rate at which work is done by the tail movements on the surrounding fluid). Small amplitude heaving and pitching motions of a finite flat-plate wing of general planform with a rounded leading edge and a sharp trailing edge are considered. This is a generalization of Chopra's (1974) work on model rectangular tails. This motion characterizes vertical oscillations of the horizontal tail flukes of some cetacean mammals. The same oscillations, turned through a right angle to become horizontal motions of side-slip and yaw, characterize the caudal fins of certain fast-swimming fishes; viz. wahoo, tunny, wavyback skipjack, etc., from the Percomorphi and whale shark, porbeagle, etc., from the Selachii. Davies’ (1963, 1976) method of finding the loading distribution on the wing and generalized force coefficients, through approximate solution of an integral equation relating the loading and the upwash (lifting-surface theory), is used to find the total thrust and the rate of working of the tail, which in turn specify the hydromechanical swimming performance of the animals. The physical parameters concerned are the tail aspect ratio ((span)2/planform area), the reduced frequency (angular frequency x typical length/forward speed), the feathering parameter (the ratio of the tail slope to the slope of the path of the pitching axis), the position of the pitching axis, and the curved shapes of the leading and trailing edges. The variation of the thrust and the propulsive efficiency with these parameters has been discussed to indicate the optimum shape of the tail. It is found that, compared with a rectangular tail, a curved leading edge as in lunate tails gives a reduced thrust contribution from the leading-edge suction for the same total thrust; however, a sweep angle of the leading edge exceeding about 30° leads to a marked reduction of efficiency. Another implication of the present analysis is that no negative work is involved in the actual oscillation of the tail.The present results are used to obtain an estimate of the drag coefficient for the motion of the animals, based on observed data and the computed thrust. The results show some evidence of differences between the CD's for cetacean mammals and scombroid fish respectively. Some discussion of this difference is also given.

Large amplitude lunate-tail theory of fish locomotion

The two-dimensional theory of lunate-tail propulsion is extended to motions of arbitrary amplitude, regular or irregular, so that an accurate comparison may be made with the actual lunate-tail propulsion of scombroid fishes and cetacean mammals. There is no restriction at all on the amplitude of motion but the tail's angle of attack relative to its instantaneous path through the water is assumed to remain small. The theory is applied to the regular finite amplitude motion of a thin aerofoil with a rounded leading edge to take advantage of the suction force and a sharp trailing edge to ensure smooth tangential flow past the rear tip. This can represent the vertical motions of the horizontal lunate tails of large aspect ratio with which cetacean mammals propel themselves or the horizontal undulations of the vertical lunate tails of certain fast fishes. The dependence of the thrust, the hydromechanical propulsive efficiency and the energy wasted in churning up the eddying wake on the reduced frequency, the angle of attack and the amplitude of motion is exhibited.

Hydromechanics of lunate-tail swimming propulsion

This paper investigates the non-uniform motion of a thin plate of finite aspect ratio, with a rounded leading edge and sharp trailing edge, executing heaving and pitching oscillations at zero mean lift. Such vertical motions characterize the horizontal lunate tails with which cetacean mammals propel themselves, and the same motions, turned through 90° to become horizontal motions of sideslip and yaw, characterize the vertical lunate tails of certain fast-swimming fishes. An oscillating vortex sheet consisting of streamwise and spanwise components is shed to trail behind the body and it is this additional feature of the streamwise component resulting from the finiteness of the plate that makes this study a generalization of the two-dimensional treatment of lunate-tail propulsion by Lighthill (1970). The forward thrust, the power required, the energy imparted to the wake and the hydromechanical propulsive efficiency are determined for this general motion as functions of the physical parameters defining the problem: namely the aspect ratio, the reduced frequency, the feathering parameter and the position of the pitching axis. The dependence of the thrust coefficient and propulsive efficiency on these physical parameters, for the complete range of variation consistent with the assumptions of the problem, has been depicted graphically.