Batoid locomotion

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Batoids are a superorder of cartilaginous fish consisting of skates, rays and other fish all characterized by dorsoventrally flattened bodies and large pectoral fins fused to the head. This distinctive morphology has resulted in several unique forms of locomotion. Most Batoids exhibit median paired fin swimming, utilizing their enlarged pectoral fins. Batoids that exhibit median paired fin swimming fall somewhere along a spectrum of swimming modes from mobuliform to rajiform based on the number of waves present on their fin at once. Of the four orders of Batoidae this holds truest for the Myliobatiformes (rays) and the Rajiformes (skates). The two other orders: Rhinopristiformes and Torpediniformes exhibit a greater degree of body caudal fin swimming. [1] [2]

Contents

Mobuliform swimming

A manta ray executing several different turns. By varying the shape of its fins asymmetrically it is able to be quite maneuverable for its size and rigidity.

Mobuliform swimming is common in pelagic Myliobatiformes species such as manta rays and is characterized by a flapping motion of the pectoral fins. It is very similar in appearance to flight in birds. Batoids that utilize mobuliform swimming can be identified by their high aspect ratios, thicker pectoral fins that taper to a point and a lateral profile that resembles a hydrofoil. They are highly efficient open water swimmers capable of traversing great distances at high speeds. [2]

The pectoral fins of a mobuliform swimming ray experience a spanwise dorsoventral deformation that is highest at the tip and a chord-wise traveling wave. [3] Kinematically mobuliform swimming consists of low frequency, high amplitude fin flapping with less than one waveform present on the fin at a time. In order to increase speed pelagic rays will increase the frequency of pectoral flaps. [4]

Rays are at a disadvantage compared to other fish when it comes to maneuverability. Their rigid body gives them a high moment of inertia and their dorsoventrally flattened shape makes it difficult to maintain turns because they are unable to provide the lateral forces necessary to prevent slip. [5] Banking during a turn has been exhibited across both types of median paired fin swimming and it allows them to compensate for the lack of control surface that they would have in an unbanked turn. Mobuliform swimmers tend to be just as maneuverable as rajiform swimmers, even though their turning mechanics are different; the former move with gliding turns while the latter move through asymmetrical undulations of the fins. [6] However, some species like the Pelagic Stingray are more maneuverable because they are able to reverse the wave along their fins and even swim backwards. [7]

In a pelagic environment rays will encounter surface waves. Experimental interactions with incoming waves for Cownose rays has shown that rays will cease swimming and form a positive dihedral with their pectoral fins allowing them to maintain their position in the water column. When travelling in the same direction of a wave it has been shown that they will increase their speed while reducing the amplitude of their fins which indicates that they may use travelling waves to increase their swimming efficiency. [8]

Rajiform swimming

An example of a rajiform myliobatoid. Notice that the movement is restricted to the distal part of the fin unlike mobuliform swimming.

Rajiform swimmers move by undulating the distal parts of their pectoral fins with multiple waveforms present on the fin at a time. This mode of swimming is utilized by demersal Batoids, which includes skates as well as some rays. They share a common morphology of a low aspect ratio and thin pelvic fins. They are slower than mobuliform swimmers but they are some of the most metabolically efficient elasmobranch swimmers at slow speeds. [9]

There are differences between the ways skates and benthic rays utilize rajiform swimming. Rajiformes do not always utilize rajiform swimming. They have a second set of pelvic fins called crura on the ventral side near the base of their tale that they use to in tandem to push along the substrate while their disk remains inactive. This style of locomotion is known as punting and is very similar to walking as the force appears to be generated from direct contact with the ground. They are able to execute asynchronous movements with their crura to make turns which negates the need to bank during turns, which may provide stealth benefits in addition to the reduced water movement. From muscle fiber analysis it appears that punting may be a primary mode of transportation at low speeds (about 1/3 Body lengths per second) in some skates and rajiform locomotion may be used when for specific situations. [10] Benthic rays rely entirely on rajiform locomotion. Another difference between the two is the role of the tail. Skates have larger tails with fins on them and they use them during turns. [6] The tails of rays appear to serve no function in swimming. Some rays, known as stingrays have a venomous barb on their tail that they whip around to defend themselves.

The distribution of pectoral thin thickness is such that rajiform swimmers benefit passively from hydrodynamic interaction between the substrate and their fins. [11] As such swimming away from the substrate for extended periods is unsustainable. [9] The thickness of the pelvic fins is highest at the anterior part of fin and lowest at the distal parts of the fin and the posterior fin, generally less than a millimeter. These thinner areas deform passively at normal speeds and must be kept rigid at higher speeds serving to limit maximum sustainable speed in rajiform swimmers. [12]

Body caudal fin swimming

Torpediniformes

pacific electric ray (torpedo californica) Pacific Electric Ray (torpedo californica).jpg
pacific electric ray (torpedo californica)

The majority of electric rays have a distinctive style of low speed swimming that consists of periodically moving up in the water column then gliding back down. Unlike Rajiformes and Myliobatiformes their propulsion comes solely from the movement of their caudal fin, which is much more developed than in skates and rays. The disc portion of their bodies is used to increase their efficiency during the gliding portion of their swimming. [13]

Rhinopristiformes

An example of a Rhinopristiform, the guitarfish relies on undulation of its caudal fin for propulsion

Rhinopristiformes are an intermediate group between sharks and rays. There has been little study into their swimming characteristics but it can be assumed from their morphological similarity to sharks that they rely primarily on body caudal fin swimming and the pectoral fins do not generate thrust.

Batoid inspired designs

Batoids have certain characteristics that would be desirable in an underwater unmanned vehicle. The nature of their movement makes them stable platforms to carry payloads. They tend to be incredibly efficient swimmers many pelagic ray species and even some benthic species undertake very long yearly migrations. Pelagic species tend to be more efficient high speed swimmers while benthic ones are efficient at lower speeds. Many Benthic rays have adapted to be incredibly stealthy, they have a low profile and create very little disturbance when they move. They have the potential to generate large thrust; this is what allows giant manta rays to completely clear the surface of the water. [14] The variations in performance capabilities of each species lead to the development of a variety of different biomimetic automated underwater vehicles (BAUVs). There are a multitude of designs based on pelagic and benthic batoids, there are even some based on more obscure aspects of batoid swimming such as one based on the unique body caudal fin propulsion of the electric ray or another that utilizes the punting seen in skates. One thing that really sets the performance of the biological and artificial versions apart is the nuanced flexibility and actuation of the disc. Different parts of the disc are considerably more flexible than others and some parts are designed to passively deform. It is especially hard to mimic the mixture of passive and active interactions of the disc of a rajiform and the ground. The complex actuation of the wings has been mimicked successfully through a variety of means including tensegrity structures, electroactive polymers, and fluid muscles. However, these technologies are not developed to the point where they can fully imitate actual muscles. To this aim, actual muscles have been used in a tissue engineered ray less than 20 mm in diameter. It was produced using rat myocardial cells that mimicked the pattern of a rajiform swimmer through the use of patterned muscle junctions. [15]

Related Research Articles

<span class="mw-page-title-main">Fin</span> Thin component or appendage attached to a larger body or structure

A fin is a thin component or appendage attached to a larger body or structure. Fins typically function as foils that produce lift or thrust, or provide the ability to steer or stabilize motion while traveling in water, air, or other fluids. Fins are also used to increase surface areas for heat transfer purposes, or simply as ornamentation.

<span class="mw-page-title-main">Stingray</span> Suborder of fishes

Stingrays are a group of sea rays, which are cartilaginous fish related to sharks. They are classified in the suborder Myliobatoidei of the order Myliobatiformes and consist of eight families: Hexatrygonidae, Plesiobatidae, Urolophidae (stingarees), Urotrygonidae, Dasyatidae, Potamotrygonidae, Gymnuridae and Myliobatidae . There are about 220 known stingray species organized into 29 genera.

<span class="mw-page-title-main">Myliobatiformes</span> Order of cartilaginous fishes

Myliobatiformes is one of the four orders of batoids, cartilaginous fishes related to sharks. They were formerly included in the order Rajiformes, but more recent phylogenetic studies have shown the myliobatiforms to be a monophyletic group, and its more derived members evolved their highly flattened shapes independently of the skates.

<span class="mw-page-title-main">Gymnotiformes</span> Order of bony fishes

The Gymnotiformes are an order of teleost bony fishes commonly known as Neotropical knifefish or South American knifefish. They have long bodies and swim using undulations of their elongated anal fin. Found almost exclusively in fresh water, these mostly nocturnal fish are capable of producing electric fields to detect prey, for navigation, communication, and, in the case of the electric eel, attack and defense. A few species are familiar to the aquarium trade, such as the black ghost knifefish, the glass knifefish, and the banded knifefish.

<span class="mw-page-title-main">Skate (fish)</span> Family of fishes

Skates are cartilaginous fish belonging to the family Rajidae in the superorder Batoidea of rays. More than 150 species have been described, in 17 genera. Softnose skates and pygmy skates were previously treated as subfamilies of Rajidae, but are now considered as distinct families. Alternatively, the name "skate" is used to refer to the entire order of Rajiformes.

<span class="mw-page-title-main">Bluegill</span> Species of fish

The bluegill, sometimes referred to as "bream," "brim," "sunny," or "copper nose" as is common in Texas, is a species of North American freshwater fish, native to and commonly found in streams, rivers, lakes, ponds and wetlands east of the Rocky Mountains. It is the type species of the genus Lepomis, from the family Centrarchidae in the order Perciformes.

<span class="mw-page-title-main">Animal locomotion</span> Self-propulsion by an animal

Animal locomotion, in ethology, is any of a variety of methods that animals use to move from one place to another. Some modes of locomotion are (initially) self-propelled, e.g., running, swimming, jumping, flying, hopping, soaring and gliding. There are also many animal species that depend on their environment for transportation, a type of mobility called passive locomotion, e.g., sailing, kiting (spiders), rolling or riding other animals (phoresis).

<span class="mw-page-title-main">Fish locomotion</span> Ways that fish move around

Fish locomotion is the various types of animal locomotion used by fish, principally by swimming. This is achieved in different groups of fish by a variety of mechanisms of propulsion, most often by wave-like lateral flexions of the fish's body and tail in the water, and in various specialised fish by motions of the fins. The major forms of locomotion in fish are:

<span class="mw-page-title-main">Rajiformes</span> Order of fishes in the superorder Batoidea

Rajiformes is one of the four orders in the superorder Batoidea, flattened cartilaginous fishes related to sharks. Rajiforms are distinguished by the presence of greatly enlarged pectoral fins, which reach as far forward as the sides of the head, with a generally flattened body. The undulatory pectoral fin motion diagnostic to this taxon is known as rajiform locomotion. The eyes and spiracles are located on the upper surface of the head and the gill slits are on the underside of the body. Most species give birth to live young, although some lay eggs enclosed in a horny capsule.

<span class="mw-page-title-main">Urolophidae</span> Family of cartilaginous fishes

The Urolophidae are a family of rays in the order Myliobatiformes, commonly known as stingarees or round stingrays. This family formerly included the genera Urobatis and Urotrygon of the Americas, which are presently recognized as forming their own family Urotrygonidae. Stingarees are found in the Indo-Pacific region, with the greatest diversity off Australia. They are sluggish, bottom-dwelling fish that have been recorded from shallow waters close to shore to deep waters over the upper continental slope. Measuring between 15 and 80 cm long, these rays have oval to diamond-shaped pectoral fin discs and relatively short tails that terminate in leaf-shaped caudal fins, and may also have small dorsal fins and lateral skin folds. Most are smooth-skinned, and some have ornate dorsal color patterns.

<span class="mw-page-title-main">Southern stingray</span> Species of fish

The southern stingray is a whiptail stingray found in tropical and subtropical waters of the Western Atlantic Ocean from New Jersey to southern Brazil. It has a flat, diamond-shaped disc, with a mud brown, olive, and grey dorsal surface and white underbelly. The barb on its tail is serrated and covered in a venomous mucus, used for self-defense.

<span class="mw-page-title-main">Pelagic stingray</span> Species of cartilaginous fish

The pelagic stingray is a species of stingray in the family Dasyatidae, and the sole member of its genus. It is characterized by the wedge-like shape of its pectoral fin disc, which is much wider than long, as well as by the pointed teeth in both sexes, whip-like tail with extremely long tail spine, and uniform violet to blue-green coloration. It generally reaches 59 cm (23 in) in width. The pelagic stingray has a worldwide distribution in waters warmer than 19 °C (66 °F), and migrates seasonally to spend the summer closer to the continental shelf and at higher latitudes. The only stingray that almost exclusively inhabits the open ocean, this species is typically found in surface waters down to a depth of 100 m (330 ft). As a consequence of its midwater habits, its swimming style has evolved to feature more of a flapping motion of the pectoral fins, as opposed to the disc margin undulations used by other, bottom-dwelling stingrays.

<span class="mw-page-title-main">Smooth butterfly ray</span> Species of fish

The smooth butterfly ray is a species of cartilaginous fish in the family Gymnuridae. It is a member of the order Myliobatiformes, which contains 10 total families. Its natural habitats are shallow seas, subtidal aquatic beds, estuarine waters, and coastal saline lagoons. Its common name is derived from its compressed body, pectoral fins that are wider than their length, and overall diamond shape.

<span class="mw-page-title-main">Aquatic locomotion</span>

Aquatic locomotion or swimming is biologically propelled motion through a liquid medium. The simplest propulsive systems are composed of cilia and flagella. Swimming has evolved a number of times in a range of organisms including arthropods, fish, molluscs, amphibians, reptiles, birds, and mammals.

<span class="mw-page-title-main">Fin and flipper locomotion</span>

Fin and flipper locomotion occurs mostly in aquatic locomotion, and rarely in terrestrial locomotion. From the three common states of matter — gas, liquid and solid, these appendages are adapted for liquids, mostly fresh or saltwater and used in locomotion, steering and balancing of the body. Locomotion is important in order to escape predators, acquire food, find mates and bury for shelter, nest or food. Aquatic locomotion consists of swimming, whereas terrestrial locomotion encompasses walking, 'crutching', jumping, digging as well as covering. Some animals such as sea turtles and mudskippers use these two environments for different purposes, for example using the land for nesting, and the sea to hunt for food.

<span class="mw-page-title-main">Batoidea</span> Superorder of cartilaginous fishes

Batoidea is a superorder of cartilaginous fishes, commonly known as rays. They and their close relatives, the sharks, comprise the subclass Elasmobranchii. Rays are the largest group of cartilaginous fishes, with well over 600 species in 26 families. Rays are distinguished by their flattened bodies, enlarged pectoral fins that are fused to the head, and gill slits that are placed on their ventral surfaces.

<span class="mw-page-title-main">Tradeoffs for locomotion in air and water</span> Comparison of swimming and flying, evolution and biophysics

Certain species of fish and birds are able to locomote in both air and water, two fluid media with very different properties. A fluid is a particular phase of matter that deforms under shear stresses and includes any type of liquid or gas. Because fluids are easily deformable and move in response to applied forces, efficiently locomoting in a fluid medium presents unique challenges. Specific morphological characteristics are therefore required in animal species that primarily depend on fluidic locomotion. Because the properties of air and water are so different, swimming and flying have very disparate morphological requirements. As a result, despite the large diversity of animals that are capable of flight or swimming, only a limited number of these species have mastered the ability to both fly and swim. These species demonstrate distinct morphological and behavioral tradeoffs associated with transitioning from air to water and water to air.

<span class="mw-page-title-main">Clearnose skate</span> Species of cartilaginous fish

The clearnose skate is a species of cartilaginous fish in the family Rajidae. R. eglanteria is also known by other common names such as the brier skate and summer skate. Clearnose skates are easily identified by the translucent patches on either side of their snouts and their mottled dorsal surface. They are found along the Atlantic and Gulf coasts of the United States in shallow waters of the continental shelf.

<span class="mw-page-title-main">Fish fin</span> Bony skin-covered spines or rays protruding from the body of a fish

Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of Actinopterygii and Chondrichthyes fishes. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as seen in sharks. Apart from the tail or caudal fin, fish fins have no direct connection with the spine and are supported only by muscles. Their principal function is to help the fish swim.

<span class="mw-page-title-main">Pelvic fin</span> Paired fins located on the ventral surface of fish

Pelvic fins or ventral fins are paired fins located on the ventral (belly) surface of fish, and are the lower of the only two sets of paired fins. The pelvic fins are homologous to the hindlimbs of tetrapods, which evolved from lobe-finned fish during the Middle Devonian.

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