Highly simplified phylogeny of ray-finned fishes with a chondrichthyian outgroup clade to show the major pattern of caudal fin evolution from the primitive heterocercal condition to the derived homocercal configuration. Caudal fin cells. Inappropriate injection positions will cause a low immunization rate and even death. Two alternative thrust directions are shown for the heterocercal tail; one based on the classical model of heterocercal tail lift (upper arrow), and the other on the model of Thomson (1976, lower arrow), in which thrust is directed toward the center of mass of the fish. The reactive force on the fish (FR) is thus oriented anterodorsally. The evolution of the modern design, with a symmetrical tail and highly manoeuvrable body fins, was only possible after the evolution of the swim bladder. See answer (1) Best Answer. Pell, and S.A. Wainwright. Homocercal tail kinematics in a bluegill, Lepomis macrochirus, swimming at 1.2 lengths/sec. Bookshelf Z-dimension excursions (upper panel) of two points on the tail and the three-dimensional angles of two tail triangles with the XZ plane. This review examines the caudal fin throughout the main lineages of fish evolution, and highlights where changes in shape and usage have occurred. Careers. Homocercal tails are the most common caudal fin type in fish but can come in many symmetrical shapes. Is differential motion of the dorsal and ventral lobe possibly a passive consequence of internal skeletal asymmetries or is it actively generated? 3. While quantifying the three-dimensional motion of the caudal fin is one critical component of understanding caudal fin function, it is also necessary to evaluate the impact that movement of the fin has on the fluid. Given this orientation, the position of the three points in a two-dimensional view is shown on the right. In homocercal tails, the vertebral column typically terminates near the base of the skeletal elements supporting the tail (hypural bones in teleosts), and although the internal caudal skeleton is not completely dorsoventrally symmetrical, the dorsal and ventral lobes of the tail are nearly equivalent in area and composition. What is the main function of caudal fins? The tail fin, also called the caudal fin, helps propels fish forward. If the homocercal tail were functioning as a homogeneous flat vertical plate and generating a reactive force directly forward with no Y component (as the classical hypothesis predicts), the dorsal and ventral tail lobes should both maintain a 90 angle to the horizontal throughout the tail beat cycle. A homocercal tail is contrasted with a heterocercal tail which has unequal lobes. What is the function of dorsal fin? This result, combined with DPIV flow visualization data, suggest a new hypothesis for the function of the homocercal tail: the homocercal tail generates tilted and linked vortex rings with a central jet inclined posteroventrally, producing an anterodorsal reactive force on the body which generates lift and torque in the manner expected of a heterocercal tail. The basic classification of fin shape dates from Louis Agassiz in 1833 who proposed the terms heterocercal (for externally asymmetrical tails with larger dorsal lobes containing the terminal extension of the vertebral column or notochord) and homocercal (for tails which are externally symmetrical and have equal-sized upper and lower lobes). An alternative model in which the shark tail produces a reactive force that acts through the center of mass is not supported. In order to record three-dimensional data on caudal fin movements during steady locomotion, I have used the experimental design illustrated in Figure 5. The functional patterns described above for bluegill are thus likely to be widespread among teleosts, although only a thorough comparative study of caudal function in fishes will reveal the extent of functional diversity. 13:476-493. Of the main vertebrate groups, fishes have occupied the planet for the longest Is it okay for an 18 year old to date a 16 year old in Florida? If the caudal peduncle is narrow and the caudal fin is forked, the fish is a fast swimmer. The central jet through the ring has a slight ventral inclination depicted diagrammatically by the large colored arrow. Although there are four types, Regular exam updates, QnA, Predictors, College Applications & E-books now on your Mobile. Lauder, G.V., C. Connon, and D. Dunn-Rankin. Caudal fin: Also known as the tail fin, the caudal fin propels the fish through the water. Because of the prominence of the caudal fin in early vertebrate fossils, its importance in locomotion, and the diversity of tail shapes, nearly all textbooks of vertebrate paleontology, anatomy, and ichthyology discuss the evolution of the tail. DuBois, A.B., G.A. The caudal fin is divided into two lobes: a larger dorsal lobe and a smaller ventral lobe. The Spines, when held stiff, make small fish much harder to swallow. Therefore, the early fish evolved an asymmetrical tail which supplies forward momentum with the top lobe and lift with the smaller bottom lobe (which, not being stiff, flexes in counterpoint to the upper lobe). This review examines the caudal fin throughout the main lineages of fish evolution, and highlights where changes in shape and usage have occurred, creating a coherent picture of caudAL fin evolution. The current literature contains several hypotheses about the function of caudal fins of different shape, and much of this discussion has focused on the difference between heterocercal and homocercal tails (Alexander, 1965; Aleev, 1969; Magnuson, 1970; Thomson, 1971, 1976; Hopson, 1974; Thomson and Simanek, 1977; Webb and Smith, 1980). Who is the blond woman in Jon Secada's Just Another Day video? It is the primary Whereas fresh water has a density of 1.0 and saltwater of about 1.026. Measurement of fluid motion in the wake of the tail of a swimming sturgeon (Liao and Lauder, 2000) reveals two counter-rotating centers of vorticity (in the XY plane) which reflect a section through a vortex ring shed by the beating tail. WebAnal Fin: is located on the ventral side of the fish, anterior to the caudal fin. George V. Lauder, Function of the Caudal Fin During Locomotion in Fishes: Kinematics, Flow Visualization, and Evolutionary Patterns, American Zoologist, Volume 40, Issue 1, February 2000, Pages 101122, https://doi.org/10.1093/icb/40.1.101. Not all fish have anal fins, but they can also be very tiny and hard to see. These X and Z component patterns are similar to those seen for sharks as expected given oscillatory tail movement and the necessity for thrust production. Note the projection of the triangle onto the XZ plane and the relative positions of the vertices on the Z-axis. WebEach fin on a fish is designed to perform a specific function: Adipose fin. Reddish color indicates fluid rotation in a counter-clockwise direction, while blue colors reflects clockwise fluid rotation; green colors indicate minimal fluid rotation. Of course, the fish has to keep moving if it takes a rest it will still sink. The cookie is used to store the user consent for the cookies in the category "Analytics". The caudal fin, or tail fin, is located at the end of the fish and provides the power that propels the fish forward, like a motor. 2). Adipose fin development and its relation to the evolutionary origins of median fins. The arrows in Figure 8 show the direction of dorsal and ventral tail movement, and it is clear that for much of the tail beat the dorsal and ventral lobes of the tail are moving in opposite directions. 1A]), to the better known caudal morphologies in ostracoderms (Fig. The video image below shows a sample image obtained from a bluegill sunfish (Lepomis macrochirus) swimming at 1.2 lengths/sec. Analysis proceeds by choosing pairs of video images (separated in time by 4 ms) that capture flow in the wake behind the tail. Schematic representation of the DPIV (digital particle image velocimetry) technique for the study of fluid flow. Caudal (Tail) Fin. 11): the homocercal tail of bluegill thus functions asymmetrically during steady swimming. The lateral line is a sensory system that enables fish 2 What are the different types of caudal fins? it is called the Caudal Fin The fin is the "Caudal Fin" and the tail part of the body the Currently, traditional visual 2022 Aug 24;5(1):863. doi: 10.1038/s42003-022-03730-0. We endeavor to keep you informed and help you choose the right Career path. Functional analysis of the homocercal tail of bluegill shows that the dorsal and ventral lobes do not function symmetrically as expected. The Question containing Inaapropriate or Abusive Words, Question lacks the basic details making it difficult to answer, Topic Tagged to the Question are not relevant to Question, Question drives traffic to external sites for promotional or commercial purposes, The Answer containing Inaapropriate or Abusive Words, Answer drives traffic to external sites for promotional or commercial purposes. Indeed, we lack even basic kinematic data on the movement of the caudal fin during steady swimming, and have even less information on the effect of different tail shapes on patterns of fluid flow and thrust production. The size and arrangement the fin determines how quickly and in what manner a fish swims. Given a 20*20 matrix of areas, a regularly spaced array of 400 velocity vectors is obtained that provides a quantitative estimate of flow in the light sheet at that time. WebAll fish have external appendages called fins. Like the tail functions of other vertebrate groups, the fish caudal fin can provide balance and lift, and can be used to capture prey or for defense. 1E, F, 2) despite considerable diversity in the morphology of tail shapes in fossil fishes (see Moy-Thomas and Miles, 1971). Fish pictures courtesy of S. M. McGinnis (1984, Freshwater Fishes of California. The Regents of the University of California.). The adipose fin is sort of a mystery to people who study fish. Table 1 Movement of four markers (M1M4, from dorsal to ventral) on the trailing edge of the caudal fin of bluegill sunfish (Lepomis macrochirus) during steady swimming at two speeds.*. WebScore: 4.2/5 (9 votes) . In order to facilitate repeated and accurate recognition of specific locations on the fin, fish are anesthetized prior to each experiment and small markers are glued bilaterally onto the fin. Would you like email updates of new search results? While the classical view of shark tail function was corroborated by the three-dimensional kinematic study of leopard sharks, the pattern of heterocercal tail function in sturgeon proved to be quite different. The sinus venosus receives venous blood from the different parts of the body through paired Cuvierian ducts and single hepatic vein. The reactive force from the vortex rings shed by the tail may be thus directed through the body near the center of mass. The fins of all fish are supported by fin rays, however in cartilaginous fish these fin rays are inflexible, unbranched and unsegmented. Such torques are proposed to be counteracted anteriorly by lift forces generated by the head and pectoral fins. What are the functions of fins? Movement of the optical components allows the laser light sheet to be oriented into three orthogonal planes (video cameras are also appropriately repositioned to provide an image of the light sheet), and video images are taken of the light reflected from the particles in the flow (also see Drucker and Lauder, 1999). Fluid velocity vectors (small arrows) and vorticity (colors) in a vertical plane behind the tail of a bluegill, Lepomis macrochirus, swimming at 1.6 lengths/sec. Finally, the significance of the diversity of tail designs in early vertebrates and major evolutionary patterns to tail morphology (Figs. They could now devote the whole of the caudal fin to propulsion and develop the pectoral and pelvic fins to maximise their maneuverability. WebWhat is function of caudal fin? Raffel, M., C. Willert, and J. Kompenhans. The tail of a shark consists of the caudal peduncle and the caudal fin, which provide the main source of thrust for the shark. Bar graphs have been scaled to 100% of maximum in the X-dimension. The heterocercal tail configuration shown in F is plesiomorphic for ray-finned fishes. Fin rays that comprise the ventral lobe are relatively flexible and follow the leading dorsal edge of the tail. Some species such as Greenland sharks, Lantern sharks, and Sleeper sharks lack this pair of fins. HIPAA privacy rule applies to the following. Variation in the tail, or caudal fin, is often a reflection of a fish's environment, and affects movement, predation, defense, and reproduction. The caudal fin represents a fundamental design feature of vertebrates that predates the origin of jaws and is found in both agnathans and gnathostomes. For full access to this pdf, sign in to an existing account, or purchase an annual subscription. WebWhat does a forked caudal fin mean? These movements are actively generated by the hypochordal longitudinalis muscle within the tail. The caudal fin is the main fin for propulsion to move the fish forward. The site is secure. Measurement of projected YZ planar angles (Table 1; Fig. The dorsal lobe of the tail achieves an XZ planar angle of nearly 75, significantly less than the 90 angle expected under the classical hypothesis of homocercal tail function. Positive X values reflect a vector pointing posteriorly, while a negative X-value would indicate a vector pointing anteriorly. Such novelties in form and function tend to have far-reaching evolutionary consequences. Copy. But the hypochordal longitudinalis (HL) muscle possesses a fiber axis at an appreciable angle to the horizontal (Fig. The HL muscle originates from the ventrolateral surface of the caudal skeleton and passes posterodorsally to make four tendinous insertions on the first four fin rays. Water in a flow tank is seeded with small (12 mean diameter) silver coated glass beads which reflect light from an argon-ion laser. The caudal fin is also known as tail fin or a median fin which is usually homocercal or heterocercal. Dickson, and G.V. Fins typically function as foils that produce lift or thrust, or provide the ability to steer or Why do you have to swim between the flags? The https:// ensures that you are connecting to the From this matrix of velocity vectors, fluid vorticity, momentum, circulation, and force can be calculated (Drucker and Lauder, 1999) using standard methods (Rayner, 1979; Spedding et al., 1984; Spedding and Maxworthy, 1986; Spedding, 1987). Information from both cameras together provides X, Y, and Z coordinates for points on the fins. In some species they are used to hold on to the substrate, while in others they allow the fish to walk or even to fly. The dorsal fin extends along the length of the back, the anal fin is located on the ventral side of the fish, and behind the anal opening, and the caudal fin is the tail.The dorsal fin may extend the entire length of the back, may be a fleshy ray, or quite large and supported by spines. 8600 Rockville Pike SVCV. This allows determination of precisely which portion of the tail is acting on the fluid. 9) show that the XZ angles oscillate about a mean angle of 90 during the tail beat indicating that these triangles do not maintain a consistent acute orientation relative to the horizontal plane as does the leopard shark tail. In this paper I review two experimental approaches which promise to provide new insights into the function and evolution of the caudal fin: three-dimensional kinematic analysis, and quantitative flow measurements in the wake of freely-swimming fishes using digital particle image velocimetry (DPIV). The particles are carried through the light sheet with water movement, and as the flow is disturbed by movement of the tail particles move with the flow and their reflections are captured on video. Fig. Both organizations make no representation on the accuracy of opinions expressed in this blog. The cookie is set by GDPR cookie consent to record the user consent for the cookies in the category "Functional". Previous literature has discussed many aspects of caudal fin form and function in particular taxonomic groups; however, no previous work has synthesized these studies in order to detail how the However, the Y component displays a different pattern than seen for sharks, oscillating about a horizontal orientation, whereas in the leopard shark Y movement vectors are consistently negative. The middle panel shows literature interpretations of caudal fin function (in lateral view) with the hypothesized axis of bending and direction of thrust (Affleck, 1950). The X movement vectors are consistently positive and the Z vectors oscillate negative and then positive as the tail beats from side to side. Heidi Doan assisted greatly with collection of kinematic data from bluegill. Fins can also be used for other purposes; male sharks and mosquitofish use a modified fin to deliver sperm, thresher sharks use their caudal fin to stun prey, reef stonefish have Images or drawings of the tail of other teleosts swimming by Bainbridge (1963) and Aleev (1969) suggest that the homocercal tail of a diversity of teleost fishes exhibits asymmetry during horizontal locomotion. Fish fins are thin and often diaphanous and monochromatic, making identification of specific points difficult, while quantifying motion of a clear fluid is a difficult problem of long-standing (Nakayama, 1988; Yang, 1989; Nieuwstadt, 1993; Moin and Kim, 1997). If the HL muscle is active during steady locomotion, it could cause the dorsal fin rays to lead the ventral rays during the tail beat, resulting in the kinematic pattern shown in Figures 11, 12, and 13. These cookies will be stored in your browser only with your consent. Performance cookies are used to understand and analyze the key performance indexes of the website which helps in delivering a better user experience for the visitors. This cookie is set by GDPR Cookie Consent plugin. Evolutionary analysis of swimming speed in early vertebrates challenges the 'New Head Hypothesis'. Reliance on a lateral or ventral view alone provides similarly misleading information on motion in the other planes. This cookie is set by GDPR Cookie Consent plugin. It also Scale bar is 1 cm. The higher the bar, the greater the velocity and area of the tail in that dimension. Schematic view of flow tank, mirror and video camera arrangement used to obtain three-dimensional data from swimming fish. Figure 8 shows six representative video frames, each illustrating a simultaneous lateral and posterior view of the tail. The interradialis and flexor muscles within the tail show no activity at this speed. 13). Under this hypothesis, the homocercal tail does not function in a manner consistent with the classical model, and generates lift forces and torques that must be balanced by anterior forces generated by the orientation and/or movement of the body and pectoral or pelvic fins. When you look back in life , this app would have played a huge role in laying the foundation of your career decisions. What effect does this asymmetrical function of the dorsal and ventral tail lobes have on patterns of water flow in the wake? For example, Romer (1966, p. 5) includes a discussion of caudal fin structure in his introductory chapter on basic vertebrate features, and similar analyses can be found in Hildebrand (1974), Carroll (1988), Pough et al. Oxford University Press is a department of the University of Oxford. Functional cookies help to perform certain functionalities like sharing the content of the website on social media platforms, collect feedbacks, and other third-party features. Lauder. Collette, and D.E. Freitas R, Gmez-Skarmeta JL, Rodrigues PN. Function of the Caudal Fin During Locomotion in Fishes: Kinematics, Flow Visualization, and Evolutionary Patterns1 George V. Lauder2 Department of Ecology and Evolutionary Biology, What are annual and biennial types of plants? The caudal fin, or tail fin, propels a fish through the water and enables it to swim. Plot of the orientation of line segments in the YZ plane formed by posterior tail markers in bluegill swimming at 1.2 lengths/sec. The cookie is set by the GDPR Cookie Consent plugin and is used to store whether or not user has consented to the use of cookies. The Perch has two dorsal fins, on the anterior side and one on the posterior end. Oh - and he wrote this website. Your report is submitted, the author will be informed about it. J. Exp. The experimental arrangement is as shown in Figure 5 with the addition of a laser and light sheet extending into the flow tank. If the caudal peduncle is narrow and the caudal fin is forked, the fish is a fast swimmer. The functional significance of. The surface of the dorsal lobe also achieves a significantly acute angle to the horizontal plane suggesting that the homocercal tail of bluegill generates lift during steady swimming. The fins of cartilaginous fish also differ in their basic internal anatomy from those of bony fish. Fig. Does pastor ayo oritsejiafor have biological children? Shadwick, R., J. Steffensen, S. Katz, and T. Knower. There is considerable diversity of tail shape within the teleost fishes. 2. The caudal or tail fin in fish and pigeon is used to increase the velocity. Note that for most of the tail beat, the orientation of these two triangular elements is greater than 90 indicating that the tail is moving in accordance with the classical model of heterocercal tail function. 2022 Aug;251(8):1306-1321. doi: 10.1002/dvdy.475. 13. The caudal fin, or tail, of a fish is the only fin to be connected to the vertebral column. Your one-stop Counselling package for JEE Main, JEE Advanced and BITSAT, Your one-stop Counselling package for NEET, AIIMS and JIPMER. Image Credits:- Guppy caudal fin by D3LL license CC BY-SA 3.0. They are often found lower down on the body in evolutionarily older fish species and higher upon the body in more modern forms. Wind tunnel tests on a model of Mustelus canis (Mitchill). They are round in cross-section, stiff and unsegmented. Pough, F.H., J.B. Heiser, and W.N. Rather, the dorsal lobe undergoes greater lateral excursions and moves at higher velocities than the ventral lobe. The shape of the caudal The monochromatic sturgeon tail has been bilaterally marked with small white markers at four locations to facilitate digitizing three dimensional coordinates of tail elements. Collaborative research with Cheryl Wilga greatly increased my understanding of sturgeon locomotor function, and work with Alice Gibb and Kathy Dickson on mackerel tail function assisted in formulating my ideas on the function of homocercal tails. Ventral fins have a primary function: to assist the fish in swimming. 17) which will generate a torque about the center of mass. It furthers the University's objective of excellence in research, scholarship, and education by publishing worldwide, This PDF is available to Subscribers Only. Commentdocument.getElementById("comment").setAttribute( "id", "a6ee664f170cf451b99629bc5314e04b" );document.getElementById("ebd2ec4c6f").setAttribute( "id", "comment" ); Gordon is an ecologist with two degrees from Exeter University. The caudal fin, or tail fin, is located at the end of a fish and provides the power to move a fish forward. What does the caudal fin do to help a fish? The caudal peduncle is where the tail is attached to the body. As noted above, there are two alternative views of heterocercal tail function in sharks, and three-dimensional kinematic data are needed to distinguish between the two models. This cookie is set by GDPR Cookie Consent plugin. In order to test the generality of the conclusions described above for heterocercal tails in taxa other than sharks, I examined the kinematics and fluid flow patterns around the tail of swimming sturgeon Acipenser transmontanus. Caudal fins come in many shapes and sizes; form often follows function, Killer Whales Fin: Dimensions, Strength, Size, Uses, And Facts Animal Queries, [] fins. In general, caudal fin acts like a propeller while the caudal peduncle functions as a motor. Most sharks have heterocercal caudal fins, meaning that the backbone extends into the (usually longer) upper lobe. Without fins, a fish would just be a fancy worm. Funke, R.M. Westneat, M.W., W. Hoese, C.A. The classical model predicts that the XZ angle of tail triangles should be greater than 90 as the tail moves toward increasing Z values indicating that the triangular surfaces are oriented in a manner predicted to force water posteroventrally and hence creating a reactive lift force on the tail (Fig. Epub 2013 Sep 19. 3) gives rise to lift forces perpendicular to the horizontal and torque about the center of mass that tend to pitch the head of the fish ventrally. 12) shows that the dorsal lobe of the tail achieves a significantly acute angle to the horizontal while the ventral lobe is more vertically oriented but nonetheless still substantially acute. These are paired fins. By examining the projection of the triangle on the XZ plane and the locations of the vertices on the Z axis, it is possible to see that this triangle has been positioned so that it forms an acute angle to the XZ plane; that is, it is inclined toward increasing Z values and vertex three leads the triangle as it moves toward the XY plane. Early vertebrates show a considerable diversity of tail shapes, ranging from the forked-tail of agnathan thelodonts (Wilson and Caldwell, 1993, [Fig. The homocercal bluegill tail is thus moving in a manner indicating that lift forces may be generated and that the reactive force on the body is not horizontal in orientation. These experimental data also indicate that the function of homocercal tails is considerably more complex than previously appreciated. 3 The 40-centimetre soft robot doesn't have solid batteries - instead it is propelled by a dual-function fluid that stores energy and moves the fish's fins. One of the most prominent characteristics of early vertebrates is the elongate caudal fin bearing fin rays. Caudal fin, Dorsal fin, Pectoral fin, Ventral fin, and the It is used to identify the fish species. Note that while the ventral lobe maintains a 3D angle near 90 during the tail beat, the dorsal lobe becomes significantly acute (less than 90) indicating that dorsal lobe of the tail is moving at an angle predicted to generate lift forces. One source of error in two-dimensional kinematic analyses. These data strongly support the hypothesis that asymmetrical function of the homocercal bluegill caudal fin is achieved actively as a result of intrinsic tail musculature. The .gov means its official. MeSH The HL is the only muscle in the tail with a fiber orientation at significant angle to the horizontal axis of the body. But the vortex rings generated by the sturgeon tail are oriented with an oblique axis and a central jet of fluid directed posteroventrally. Red fibers in myotomes of the caudal peduncle show light rhythmic bursting activity typical of locomotion at this speed, just above the transition from pectoral to caudal propulsion (Gibb et al., 1994). Variation in the tail, or caudal fin, is often a reflection of a fish's environment, and affects movement, predation, defense, and reproduction. Graphs of three-dimensional orientations of sturgeon tail triangles (Fig. The caudal fin is known commonly as the tail fin (Table 4.6). First, I will discuss two experimental approaches to the study of caudal fin function in fishes that are likely to greatly enhance our ability to quantify relevant functional attributes of fish fins. These are abilities which have greatly increased the habitats they can take advantage of and undoubtedly contributed considerably to their being the most numerous group of vertebrates on the planet. An Anal fin helps keep the fish upright and moving in a straight line. Fig. sharing sensitive information, make sure youre on a federal Analyses of variance conducted for leopard sharks swimming in this apparatus (Ferry and Lauder, 1996) and similar analyses for bluegill showed that the presence of the mirror in the flow had no significant effect on either tail beat amplitude or frequency (P > 0.27), suggesting that the mirror has little impact on the kinematics of the tail beat. Where is the dorsal fin and 111112) has succinctly summarized the classical view of heterocercal tail function by noting that the ventral lobe of the tail, being more flexible than the dorsal lobe, which contains the vertebral column, lags behind as the tail swings from side to side and passes through the water at an angle to the vertical. Motion of the tail at an angle to the vertical (Fig. Each video frame is split into a lateral portion on the left and a posterior view on the right and resulted from the experimental arrangement illustrated in Figure 5. Generally, it is a vertically expanded structure which is located at the caudal end of the body. Pectoral fins are paired fins that are normally located just behind the gills. Thomson (1976) proposed a different view of heterocercal tail function based on his observation that the ventral lobe of the tail appeared to lead the dorsal lobe in films taken behind the tail during locomotion by sharks in aquaria. The main types of Caudal fins found in bony fish are: A homocercal tail is a caudal fin composed of two lobes of equal proportion. Heterocercal tail kinematics in the sturgeon, Acipenser transmontanus. Because the vertebral column or notochord extends into the dorsal lobe of the tail in fishes with heterocercal caudal fins, this dorsal lobe has classically been believed to be stiffer and hence to lead the ventral lobe as it moves laterally during the tail beat (Fig. Superficial myotomal musculature and the interradialis muscles that interconnect fin rays have been removed. The caudal fin represents the distal region of the vertebrate axis and is the region of the body where fluid accelerated anteriorly is shed into the surrounding medium. The caudal fin is the tail fin, which when it is asymmetrical (as in cartilaginous fish) is termed Heterocercal and when it is symmetrical (as in most bony fish) it is termed Homocercal. Numerous different tail configurations (not shown in this figure) exist within the ray-finned fishes, even among basal clades (see e.g.,Moy-Thomas and Miles, 1971). HHS Vulnerability Disclosure, Help Out of these, the cookies that are categorized as necessary are stored on your browser as they are essential for the working of basic functionalities of the website. These cookies track visitors across websites and collect information to provide customized ads. Understanding the kinematics of dorsal and anal fins may elucidate how these fins are used in concert to maintain and change fish body position and yet little is known about the Bethesda, MD 20894, Web Policies Letters A to F indicate the times corresponding to the similarly labeled video frames in Figure 8. However, compared to the extensive analyses of myotomal muscle function that have been conducted over the last two decades (e.g., Bone et al., 1978; Johnsrude and Webb, 1985; Rome et al., 1988, 1993; Johnston, 1991; Jayne and Lauder, 1994b, 1995; Shadwick et al., 1998) and studies of axial musculoskeletal structure and function (Symmons, 1979; Hebrank, 1982; Long, 1992, 1995; Westneat et al., 1993; Jayne and Lauder, 1994a), comparatively little is known about how the tail of fishes functions during swimming. (1989), Kardong (1994), and Helfman et al., (1997). Copyright 2022 Pathfinder Publishing Pvt Ltd. The shape and structure of a fishs fins reflect both its lifestyle and its evolution. It also acts like a rudder to help a fish steer. But even the most ancient and primitive agnatha had a caudal fin. The dorsal lobe also makes an angle of 80 to the XZ plane indicating that mackerel tails function in a similar general manner to bluegill and may generate lift even during steady horizontal swimming. By understanding the fluid motion induced by action of the caudal fin, the forces exerted on the fluid and the direction of those forces can be estimated. Lower row: tail shape in a primitive gnathostome E. the arthrodire (Arctolepis) and a primitive ray-finned fish F. (Carboveles). By visualizing flow in two or more dimensions, vortices formed by fin movement can be reconstructed and the orthogonal components of momentum and force calculated (e.g.,Lauder et al., 1996; Drucker and Lauder, 1999; Wolfgang et al., 1999; Wilga and Lauder, 1999). External morphological symmetry is no guide to function: the symmetrical dorsal and ventral lobes of the homocercal tail may exhibit considerably different functional patterns with important consequences for the force balance on the body. 3). The similarity of heterocercal shape between the tails of the leopard shark and sturgeon is not mirrored by similarity of function. 11. Accessibility 4. The homocercal tail, in contrast, has been nearly universally held to generate a reaction force directed forward (near the center of mass) because of the vertical axis of bending and to move symmetrically with both dorsal and ventral lobes moving in synchrony (Fig. What are the different types of caudal fins? Fig. Rayner, and C.J. Helfman, G.S., B.B. The laser beam is focused via a series of lenses into a light sheet approximately 10 cm wide and 12 mm thick. If a car travels 400m in 20 seconds how fast is it going? Kinematic analysis was accomplished by swimming sturgeon at 1.2 lengths/sec in a flow tank as illustrated in Figure 5. Three-dimensional kinematic approaches and the ability to quantify fluid motion provide a previously unavailable perspective on the function of fish fins as control surfaces during swimming. The caudal fin has two lobes such as dorsal epichordal and ventral hypochordal lobe which are supported by the modified last three caudal vertebrae. This is a very different movement pattern than seen for the leopard shark tail. 4.22 B). official website and that any information you provide is encrypted The caudal fin extends from the tail moving from side to side. 1. Epub 2014 Feb 12. Findeis, and L. Grande. Location of the posterior-view mirror at least one to two body lengths posterior to the trailing edge of the tail and against the downstream flow grid (which restricts recirculatory vortices downstream from the mirror and hence their impact on flow immediately upstream from the mirror) minimizes any disturbance of the flow caused by the mirror in the region of the swimming fish. Six markers were attached bilaterally to the tail (three each to the upper and lower lobes) to allow quantification of tail surface orientation in three-dimensions. What is the function of a squid's fin? Fins: At the tail (posterior) end of the squid students will notice the two fins on the mantle. The fins allow the squid to steer themselves , help stabilize their position and propel the squid at slow speeds. The upper panel shows the XY position of the three vertices, and the lower panel the position of the vertices in the YZ plane. The anal fin is used for balance and steering. Fig. But opting out of some of these cookies may affect your browsing experience. Fox. Like human limbs, fins provide fish with balance, steering, and protection. Indeed, the caudal fin is a major component of mobility in fishes because it generates swimming power and contributes to maneuverability, thus improving control of swimming speed, agility, prey capture, and predator avoidance (Lauder, 2000;Flammang and Lauder, 2009). This website uses cookies to improve your experience while you navigate through the website. Note that the dorsal tail lobe (segment 12) makes an acute angle to the horizontal as the tail beats from left to right. This cookie is set by GDPR Cookie Consent plugin. 10. However, analysis of steady swimming in chub mackerel (Scomber japonicus) (Gibb et al., 1999) reveals a similar pattern of asymmetry with the dorsal lobe undergoing a 15% greater Z excursion than the ventral lobe. In eels and eel-like fish it may be united with the upper lobe of the caudal fin. In []. For most of the tail beat, the XZ angles are greater than 90 supporting the classical model of heterocercal tail function. They are the most variable, in terms of placement, of all a fishs fins. The function of the adipose fin is being studied by biologists. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. The most dorsal marker also has a higher lateral velocity than the ventral (Table 1) at both 1.2 and 2.2 lengths/sec. This fin is soft and fleshy, like an earlobe. What is the use of tail for fish and pigeon? In addition, the hypochordal longitudinalis muscle is found in virtually all teleost clades (Marshall, 1971; Lauder, 1989) and may be a key feature of the functional design of the teleost caudal fin. Analytical cookies are used to understand how visitors interact with the website. Rome, L.C., R.P. These methods are then applied to the function of the caudal fin during steady swimming in fishes with heterocercal and homocercal morphologies: chondrichthyians (leopard sharks) and ray-fined fishes (sturgeon and bluegill sunfish). The caudal fin, or tail fin, is located at the end of a fish and provides the power to move a fish forward. Fish caudal fin is a prominent example of biological propulsion, in which the caudal peduncle, fin rays and fin membrane together form a dynamic locomotory system. For each fish, generally, there are five main fins follows: Dorsal, Pelvic, Caudal (tail), Anal and Pectoral as shown in Fig. Fins are essential to fish. Fig. A fin is a thin component or appendage attached to a larger body or structure. What is the answer to the brain teaser T I M E ABDE? In addition, it is possible that the posterior views that formed the initial evidence for an alternative to the classical model were subject to the difficulties diagrammed in Figure 4: if only a posterior view is available, a surface may appear to be in a substantially different orientation from its actual three-dimensional position. This paper has three aims. 15. Caudal fin is the tail fin located at the end of Caudal peduncleand is used for propulsion. What countries have only 2 syllable in their name? Why are you allowed to use the coarse adjustment when you focus the low power objective lens? The area of interest in the wake (typically a 10 cm2 region, see Fig. Definition of caudal fin. 2013 Oct;55(8):687-98. doi: 10.1111/dgd.12088. The hypochordal longitudinalis muscle shows strong activity during steady swimming, but all other intrinsic tail muscles are inactive at this speed. Since the presence of fins as control surfaces in fishes is a prominent aspect of their biological design, it is at first glance surprising that so little is known about how fins move and what effect such movements have on fluid motion. 2017 Apr 27;15(1):32. doi: 10.1186/s12915-017-0370-x. The Perch uses paried pelvic fins to navigate, stop, move up and down and go backwards. Note that the dorsal marker undergoes a much greater excursion than the ventral marker. Is it better to take a shower in the morning or at night? Epub 2022 Apr 22. Before We love to hear from our readers. Some authors have noted more complex actions of homocercal caudal fins than simple symmetrical dorsal and ventral lobe motion (Bainbridge, 1963; Fierstine and Walters, 1968; Aleev, 1969; Videler, 1975), but no study to date has quantified motion of the homocercal caudal fin to examine explicitly the classical view of symmetrical function during steady horizontal locomotion. 1 What is the main function of caudal fins? Keywords. What Is The meaning of an Angel with no face? Spedding, G.R., J.M.V. An adipose fin is a small fin without any strengthening rays, that is only found in a few groups of fish such as the Myctophidae, Osmeridae and Salmonidae as well as some catfish. The dorsal lobe of the sturgeon tail does not lead during the tail beat, and the tail is extremely flexible. Are you sure you want to delete your comment? Under this model, the reaction force generated by movement of the heterocercal tail was proposed to be directed anteroventrally through the center of mass of the body (Fig. Federal government websites often end in .gov or .mil. Heterocercal tail kinematics in the leopard shark, Triakis semifasciata swimming steadily at 1.2 lengths/sec. You will probably have noticed that sharks and rays are either moving constantly, or resting on the bottom they do not hover in the water column. 3). McFarland.(eds.). Prior to swimming fish in the flow tank, individuals were anesthetized and small white markers attached to the tail in order to provide reliable and repeatable locations for digitizing. Their principal function is to help the fish swim. This type of tail is known as a heterocercal tail. I. Lauder and Jayne (1996) showed that angles of fin surfaces estimated from two-dimensional analyses can be in error by as much as 83 from the correct three-dimensional angle (and further details about 3D angle calculations can be found in that paper). If you have any questions or if you want to get in touch with us, you can find our contact details on our About Us page. The sturgeon tail behaves mechanically as an extremely flexible sheet with flexible dorsal and ventral lobes following the central tail region. Advertisement cookies are used to provide visitors with relevant ads and marketing campaigns. The XZ plane represents the horizontal or frontal plane, the XY plane the vertical or parasaggital plane, and the YZ plane the transverse section. Such torques must be counteracted by lift forces generated by the head and/or pectoral fins anterior to the center of mass. RNA-seq. Figure 6 illustrates the basic principle of DPIV as used in our experiments visualizing flow in the wake of the caudal fin. But measuring both fin and fluid motion accurately and in a time-dependent manner is a difficult proposition. 12. Caudal fins with lobes of equal size are homocercal and caudal fins with lobes of unequal size are called heterocercal and give lift to the fish. WebApart from the tail or caudal fin, fish fins have no direct connection with the spine and are supported only by muscles. Fin modules: an evolutionary perspective on appendage disparity in basal vertebrates. Temple has emerged suggesting the adipose fin acts as a precaudal sensing organ which aids a fish in navigating turbulent water. 562 COPEIA, 1980, NO. SUMMARY. Diagram of two extant ray-finned fishes, a sturgeon, Acipenser, (above) and a bluegill, Lepomis, (below) to illustrate heterocercal and homocercal tail shapes, respectively. Movement vectors were calculated according to the procedures described in Ferry and Lauder (1996) and reflect the projected area of the tail elements and the velocity of those elements (as components in each of three dimensions) for discrete time increments within the tail beat. 10). 8. This means that they help the fish turn left it right as well as back up. Broad evolutionary patterns of caudal fin structure have now been relatively well documented in fishes, and the internal anatomy of the caudal fin is a common source of characters for phylogenetic analysis (Patterson, 1968, 1973; Schultze and Arratia, 1986, 1988; Arratia, 1991). Note that the tail expands dorsoventrally (markers 1 and 4 move in opposite Y directions) and that the timing of this movement differs in the dorsal and ventral tail lobes indicating that the dorsal and ventral regions of the homocercal tail do not function similarly. Caudal fin Cavagna, and R.S. The predecessors of legs, they give a fish lift, steering capability, braking and momentum. Without fins, a fish would just be a fancy worm. Fig. WebThe dorsal fins increase the lateral surface of the body during swimming, and thereby provide stability but at the expense of increasing drag (see also. The results indicate that the ZFIN cell line provides a useful in vitro tool for study on the gene functions and cellular responses to viral infection in fish. Thanks! What is the help word of sharks lagoon's game a perfect wife? Web5: Anal fins. The caudal fin is the primary appendage which is used for locomotion in many fishes. Commun Biol. 17). It also acts like a rudder to help a fish steer. AFS merely hosts voluntary blogs as a place for members to share their personal views with other AFS members and to engage in friendly exchanges on fish topics. Heterocercal tail kinematics in the sturgeon, Acipenser transmontanus. The caudal fin Modified from Wilson and Caldwell (1993), Carroll (1988), and Romer (1966). Calculation of three-dimensional planar angles confirms these changes in tail lobe orientation (Fig. By swimming leopard sharks, Triakis semifasciata, in a flow tank with the dual camera arrangement shown in Figure 5, Ferry and Lauder (1996) were able to quantify the movement of specific points on the heterocercal tail in three-dimensions and to calculate the three-dimensional orientation of seven triangular fin elements. The caudal fin, or tail fin, propels a fish through the water and enables it to swim. There are different tail fin shapes for different species of fish. The size and arrangement the fin determines how quickly and in what manner a fish swims. It can also be an indicator of a general type of fish and the environment in which it lives. 6) is then selected and divided into a matrix of discrete smaller areas of interrogation. Ferry and Lauder (1996) presented additional evidence in support of this model in the form of dye injection near the tail which showed that the leopard shark tail directs water in a posteroventral direction, consistent with the classical model. The cookie is used to store the user consent for the cookies in the category "Performance". Pennycuick. The caudal fin also represents the most posterior region of the vertebrate axis and is the location where fluid, accelerated by movement of the body anteriorly, is shed into the surrounding medium. The ancestors of modern fish, both bony and cartilaginous, looked in terms of their fin structure much like modern dogfish and sharks. The ventral lobe expands within the first third of the tail beat while expansion of the dorsal lobe occurs in the final third (Fig. What is the function of caudal fin Class 9? Pelvic fins were originally found towards the rear of the fish (abdominal placement). The evolution of the swim bladder and the neutral buoyancy it gives the fish which possess it, set fish free. Wolfgang, M.J., J.M. A caudal fin graced the Summer 2012 cover of Eddies magazine (fws.gov/eddies). Dorsal fins are located either on the back of the fish or its top, it tail fin The shape of the caudal fin may vary in different species from rounded to pointed, notched, emarginated, truncated, etc. MHCET Law ( 5 Year L.L.B) College Predictor, Knockout JEE Main 2022 (Easy Installments), List of Media & Journalism Colleges in India, Top Medical Colleges in India accepting NEET Score, Medical Colleges in India Accepting NEET PG, Engineering and Architecture Certification Courses, Programming And Development Certification Courses, Artificial Intelligence Certification Courses, Top Hotel Management Colleges in Hyderabad, Top Hotel Management Colleges in Tamil Nadu, Top Hotel Management Colleges in Maharashtra, Diploma in Hotel Management and Catering Technology, Top Government Commerce Colleges in India, AILET Exam Analysis 2023 (Out): Difficulty Level, Expected Cut Off Marks, Previous Year Cut offs, AILET Answer Key 2023 - Solutions, Provisional Answer Key, Calculate Marks, Raise Objections, AILET Exam Date 2023: Answer Key, Cutoff, Result, AILET Application Form 2023 (Closed) - Steps to Apply Online Registration, Eligibility, Fees, AILET Admit Card 2023 Direct Link (Out) - Download Hall Ticket Here, Top MBA Colleges in India Accepting CAT Score. As such, the morphology of the caudal fin may influence the forces exerted on the fluid by a swimming fish and in turn the reaction forces experienced by the body during locomotion. Patterns and outliers that demonstrate how form and function of this appendage are intertwined can further inform hypotheses that fill critical gaps in knowledge concerning the caudal fin. Sign in and access our Clipboard, Search History, and several other advanced features are temporarily unavailable. But examination of the shape of the tail in these images reveals that changes in thickness occur which indicate that there are as yet unrecognized alterations in caudal fin shape that are not well revealed by ventral or dorsal views (e.g.,Gray, 1933, 1968; Aleev, 1969). Facey. This second camera provides a posterior (YZ) view of tail function. What do they land on when they jump off balcony in two and a half men? Fig. Median fins can also serve other purposes, like protection in the lion fish (Fig. Fish Body Parts List 1 Caudal fin 2 Anal fin 3 Pelvic fin 4 Pectoral fin 5 Gill 6 Gill cover 7 Mouth 8 Eye 9 Nostril 10 Dorsal fins What are the names of the parts of a fish? The caudal fin of leopard sharks functions in a manner consistent with the classical model of heterocercal tail function in which the caudal surface moves at an acute angle to the horizontal plane, and hence is expected to generate lift forces and torques which must be counteracted anteriorly by the body and pectoral fins. Homocercal tail kinematics in bluegill, Lepomis macrochirus. Video images of caudal fin function in the sturgeon, Acipenser transmontanus swimming steadily at 1.2 lengths/sec. The reasons for the asymmetry in the early fishes and in modern cartilaginous fish is that in these fish the fins supply all, or most of, the lift to the fishs body. Desvignes T, Robbins AE, Carey AZ, Bailon-Zambrano R, Nichols JT, Postlethwait JH, Stankunas K. Dev Dyn. Evolution. WebVentral/Pelvic fins: Ventral fish are a pair of fins that correspond to the hind limbs of quadrupeds. WebIn this article we will discuss about the function of venous system in fishes. The model proposed by Thomson (1976) for heterocercal tail function resulted in part from film images of shark tails taken in posterior view as sharks swam in large aquaria. Under this hypothesis, the homocercal tail functions in a similar manner to the classical model for the heterocercal tail: lift forces and torques are generated posteriorly. Through integration of past and present work, this review creates a coherent picture of caudal fin evolution. While analyses of locomotion on land have traditionally used force plates to quantify the forces exerted by limbs during locomotion (Cavagna, 1975; Biewener and Full, 1992), a technique allowing similar measurements has not been available until recently for the aquatic realm. Quantitative flow visualization in the wake of bluegill swimming in a flow tank at 1.6 lengths/sec (Fig. The positions of four markers on the trailing edge of the tail are shown in the YZ plane for one-half tail beat. Small reflective particles are placed in the water and light from a laser is focused into a light sheet which reflects off of individual particles and is imaged by high-speed video. They are absent or reduced in eels and eel-like fish and in some bottom-dwelling species they have become modified into clasping organs. The tail increases in height during the tail beat, although this height increase is achieved by asymmetrical movements of the dorsal and ventral tail lobes. One camera is aimed at the lateral tail surface and provides data in the XY plane, while a second, synchronized, camera is aimed at a small mirror placed in the flow downstream from the swimming fish. This orientation indicates that the reaction force on the sturgeon is directed anterodorsally during steady horizontal locomotion, and suggests a new hypothesized force balance on swimming sturgeon. The question have been saved in answer later, you can access it from your profile anytime. Webthe fish remain stationary to pick insects off plants. By the time of the placoderms, pelvic fins had also evolved. fin are all unpaired fins. Elasmobranchs (sharks) and ray-finned fishes also show considerable diversity in caudal fin morphology, and diversification in structures involved in locomotion has been a major theme in the evolution of these clades (Lauder, 1989). The technique of DPIV (digital particle image velocimetry) provides a means of quantifying fluid flow and of calculating forces exerted by fishes swimming in vivo. The caudal fin is the tail fin, located at the end of the caudal peduncle and is used for propulsion What is tale fin of a fish is called? Unlike many marine mammals with tails that use an up-and-down motion, fish generally use a side-to-side thrust of their caudal fin for propulsion. Fins typically function as foils that produce lift or thrust, or provide the ability In some species they are used to hold on to the substrate, while in others they allow the fish to walk or even to fly. In contrast, the alternative model predicts that the XZ angle will be less than 90 with the expectation that water influenced by motion of the tail will be directed posterodorsally and thus create a reaction force directed slightly ventrally through the center of mass. Mean flow velocity has been subtracted from the U (horizontal) component of the velocity vectors to better reveal changes in flow due to the tail beat; arrow in upper left gives the velocity vector scale. The relative movements of the dorsal and ventral markers did not change over the speed range of 1.2 to 2.2 lengths/sec. It is apparent from the first frame that the dorsal lobe containing the vertebral column does not lead the tail beat. The cookies is used to store the user consent for the cookies in the category "Necessary". It also acts like a rudder to help a 7. One way in which such errors can arise is shown in Figure 4 which depicts a three-dimensional space defined by X, Y, and Z axes. Kinematics of the homocercal tail in bluegill were studied by swimming fish at 1.2, 1.6, and 2.2 lengths/sec in a flow tank as illustrated in Figure 5. Anatomically, these muscles consist of dorsal and ventral flexor muscles (which often have deep and superficial components), the carinal muscles that connect the most dorsal and ventral skeletal elements of the tail to the dorsal and anal fins, and interradialis muscles (Fig. The views expressed in these posts are solely the views of the AFS member who wrote them and do not necessarily reflect the views of the American Fisheries Society (AFS), its officers, or the U.S. Geological Survey. Schematic illustration of the vortex wake behind a bluegill sunfish swimming steadily. We also use third-party cookies that help us analyze and understand how you use this website. Lateral Line: is located laterally on each side of the body. By clicking Accept All, you consent to the use of ALL the cookies. INTRODUCTION. If the fish is happy to live on the sea floor, this is not a problem. The zebrafish caudal fin is a non-muscularised dermal fold which originates from the ventral side of the larval fin and is composed of 16-18 bony rays which can be bifurcated 2.Caudal fin rays are segmented and articulated between each other by inter-segmental joints, providing flexibility to the fin 5.Rays are made up of a pair of concave Such a space may represent the working section of a flow tank, or the aquarium within which an experiment is conducted. Although additional terms are used to describe variants of these two shapes, Agassiz's terminology is retained in all modern textbooks. Representative data from two tail triangles are plotted in Figure 7 along with data showing the lateral (Z) excursion of two points on the tail. 1). A. However, if we rely on a posterior view alone, projection of the trailing edge (line segment 2-1) onto the YZ plane is inclined dorsally suggesting, erroneously, that fluid influenced by such a motion might be directed dorsally. Video frames (top, showing the posterior (YZ) view of tail motion) and electromyographic recordings (bottom) of intrinsic caudal muscles during steady swimming in bluegill (Lepomis macrochirus) at 1.2 lengths/sec. From the Symposium on The Function and Evolution of the Vertebrate Axis presented at the Annual Meeting of the Society of Integrative and Comparative Biology, 610 January 1999, at Denver, Colorado. A must visit. kudos to the team! A tail surface element oriented in such a position and moving toward the XY plane (into the page, or toward increasingly large Z-axis values) would be expected to push water posteroventrally. and transmitted securely. The main function of caudal find is to help the fish swim. 3). But if it wishes to hunt nearer the surface, then it has to keep making an effort to stay up. Sci Rep. 2019 Jan 24;9(1):512. doi: 10.1038/s41598-018-37040-5. Plots of marker Z-dimension excursions show that the dorsal lobe of the tail undergoes approximately a 50% greater lateral movement than the ventral lobe (Fig. Spines are associated with the fins of most fish. The homocercal tail represents a derived morphology (Fig. The paired fins assist with steering, stopping and hovering. If a triangle is suspended within this space to represent the tail of a fish swimming in a flow tank, then water would flow through the YZ plane parallel to the XY plane. J Exp Zool B Mol Dev Evol. What is the function of the caudal fin of a fish? The cookie is used to store the user consent for the cookies in the category "Other. Note that as expected, positive thrust is developed throughout the tail beat as indicated by the consistently positive X values, and that the Z vector magnitudes alternate between left and right sides. Fishes are the longest persisting living vertebrates and as such, display an incredible array of diversity. BMC Biol. Two attributes of fin function in fishes that have received the least attention are (1) a precise description of the motion of surface elements of the fin and (2) an analysis of the effect that fin motions have on the water. Carter (1967, pp. They are an useful means of defence for a fish because they effectively increase its size without a great increase in living costs. What is the main function of caudal fins? Fins are either single along the centerline of the fish: the dorsal fin, anal fin, and tail fin; or paired fins: the pectoral fins and ventral fins.i Pectoral fins help fish balance. However, the Y vectors unexpectedly show an alternation between negative and positive values which suggest that the tail is generating little net vertical force. Anderson, M. Grosenbaugh, D. Yue, and M. Triantafyllou. Whereas in bony fish they are branched, flexible and segmented except in the ancient coelocanths whose finrays are unbranched. fin is attached to is called the "Caudal Peduncle". New frontiers in the evolution of fin development. The right panel shows the caudal fin as a single line in posterior view as it would appear under the classical hypotheses of heterocercal and homocercal tail function. The heterocercal caudal fin is found in a diversity of fish clades and is believed to be primitive for sharks and ray-finned fishes (Figs.
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