Swimming speed performance in coral reef fishes: field validations reveal distinct functional groups

Central to our understanding of locomotion in fishes are the performance implications of using different modes of swimming. Employing a unique combination of laboratory performance trials and field observations of swimming speed, this study investigated the comparative performance of pectoral and body-caudal fin swimming within an entire assemblage of coral reef fishes (117 species 10 families). Field observations of swimming behaviour identified three primary modes: labriform (pectoral 70 spp.), subcarangiform (body-caudal 29 spp.) and chaetodontiform (augmented body-caudal 18 spp.). While representative taxa from all three modes were capable of speeds exceeding 50 cm s⁻¹ during laboratory trials, only pectoral-swimmers maintained such high speeds under field conditions. Direct comparisons revealed that pectoral-swimming species maintained field speeds at a remarkable 70% of their maximum (lab-tested) recorded speed; species using body-caudal fin propulsion maintained field speeds at around 50% of maximum. These findings highlight a profound influence of swimming mode on performance, with the relative mechanical and energetic efficiency of each swimming mode being of major importance. Combining attributes of efficiency, maneuverability and speed in one mode of propulsion, pectoral swimming appears to be a particularly versatile form of locomotion, well suited to a demersal lifestyle on coral reefs.     

On the osteology and myology of catfish pectoral girdle, with a reflection on catfish (Teleostei: Siluriformes) plesiomorphies

The configuration of the pectoral girdle bones and muscles of numerous catfishes was studied in detail and compared with that of other siluriforms, as well as of other teleosts, described in the literature. The pectoral girdle of catfishes is composed of only three bones, which probably correspond to the posttemporo-supracleithrum (posttemporal + supracleithrum), scapulo-coracoid (scapula + coracoid), and cleithrum of other teleosts. These latter two bones constitute the place of origin of the pectoral girdle muscles. Two of these muscles are related to the movements of the pectoral fin. These two muscles correspond, very likely, to the abductor superficialis and to the adductor superficialis of other teleostean fishes. In relation to the pectoral spine (thickened first pectoral fin ray), it is usually moved by three well-developed muscles, which are probably homologous with the arrector ventralis, arrector dorsalis, and abductor profundus of nonsiluriform teleosts. The morphological diversity and the plesiomorphic configuration of these muscles, as well as of the other catfish pectoral girdle structures, are discussed.     

Development of zebrafish (Danio rerio) pectoral fin musculature

During posthatching development the fins of fishes undergo striking changes in both structure and function. In this article we examine the development of the pectoral fins from larval through adult life history stages in the zebrafish (Danio rerio), describing in detail their pectoral muscle morphology. We explore the development of muscle structure as a way to interpret the fins' role in locomotion. Genetic approaches in the zebrafish model are providing new tools for examining fin development and we take advantage of transgenic lines in which fluorescent protein is expressed in specific tissues to perform detailed three-dimensional, in vivo fin imaging. The fin musculature of larval zebrafish is organized into two thin sheets of fibers, an abductor and adductor, one on each side of an endoskeletal disk. Through the juvenile stage the number of muscle fibers increases and muscle sheets cleave into distinct muscle subdivisions as fibers orient to the developing fin skeleton. By the end of the juvenile period the pectoral girdle and fin muscles have reoriented to take on the adult organization. We find that this change in morphology is associated with a switch of fin function from activity during axial locomotion in larvae to use in swim initiation and maneuvering in adults. The examination of pectoral fins of the zebrafish highlights the yet to be explored diversity of fin structure and function in subadult developmental stages. J. Morphol. (c) 2005 Wiley-Liss, Inc.     

Wave energy and swimming performance shape coral reef fish assemblages

Physical factors often have an overriding influence on the distribution patterns of organisms, and can ultimately shape the long-term structure of communities. Although distribution patterns in sessile marine organisms have frequently been attributed to functional characteristics interacting with wave-induced water motion, similar evidence for mobile organisms is lacking. Links between fin morphology and swimming performance were examined in three diverse coral reef fish families from two major evolutionary lineages. Among-habitat variation in morphology and performance was directly compared with quantitative values of wave-induced water motion from seven coral reef habitats of different depth and wave exposure on the Great Barrier Reef. Fin morphology was strongly correlated with both field and experimental swimming speeds in all three families. The range of observed swimming speeds coincided closely with the magnitude of water velocities commonly found on coral reefs. Distribution patterns in all three families displayed highly congruent relationships between fin morphology and wave-induced water motion. Our findings indicate a general functional relationship between fin morphology and swimming performance in labriform-swimming fishes, and provide quantitative evidence that wave energy may directly influence the assemblage structure of coral reef fishes through interactions with morphology and swimming performance.     

Fibre types in the locomotory muscles of an antarctic teleost,Notothenia rossii

Summary The metabolic and structural differentiation of locomotory muscles of Notothenia rossii has been investigated. In this species sustained locomotion is achieved by sculling with enlarged pectoral fins (labriform locomotion), whilst the segmental myotomal muscle is reserved for burst activity. Red, white and subepidermal fibres can be distinguished in the trunk by histochemical and ultrastructural criteria. The main pectoral muscle (m. adductor profundus) consists entirely of red fibres. These three main fibres types show differences in histochemical staining profiles, capillarization, myofibril shape and packing, and lipid and mitochondrial content. The fractional volume of mitochondria amounts to 38% for pectoral, 30% for red myotomal and 1.9% for white myotomal fibres. Enzyme activities of red pectoral muscle are consistent with a higher potential for aerobic glucose and fatty acid oxidation than for the red myotomal fibres. Mg²⁺ Ca²⁺ -myofibrillar ATPase activities are similar for red pectoral and myotomal muscles and approximately half of those white fibres. Specialisations of N. rossii muscles associated with labriform swimming and locomotion at Antarctic temperatures are discussed.     

Ecomorphology of Locomotion in Labrid Fishes

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

Kinematics, dynamics, and energetics of rowing and flapping propulsion in fishes

The shape and motion of the pectoral fins vary considerablyamong fishes that swim in the labriform mode. Pectoral fin motionin fishes is highly variable, but one conspicuous axis of thisvariation is the rowing-flapping axis. At one extreme of thisaxis, paddle-shaped fins row back and forth in a plane thatis parallel to fish motion, while at the other extreme, wing-shapedfins flap up and down in a plane that is perpendicular to fishmotion. We have used two fish, the threespine stickleback (Gasterosteusaculeatus) and the bird wrasse (Gomphosus varius), that fallnear the extremes of the rowing-flapping axis to study the dynamic,energetic, and ecological and evolutionary consequences of thiskinematic variation. Our work confirms some traditionally heldassumptions about rowing and flapping dynamics and energeticsbut reject others. A computer simulation experiment of virtualrowing and flapping appendages makes several predictions aboutdifferences in maneuvering performance and swimming energeticsbetween rowing and flapping, which, in turn, make predictionsabout the behavior and ecological distribution of fishes thatvary along the rowing-flapping axis. Both laboratory and fieldstudies of labrid swimming ability and distribution supportthese predictions.     

Botulinum toxin injections as a method for chemically denervating skeletal muscle to test functional hypotheses: a pilot study in Lepomis cyanellus

In this study, we demonstrate that botulinum toxin can be used to chemically denervate muscles to test functional hypotheses. We injected research-grade type A botulinum toxin complex into pectoral fin abductors (abductor superficialis) of green sunfish (Lepomis cyanellus) to determine whether chemical denervation would eliminate the ability of a particular muscle to contribute to overall pectoral fin movements. Reduction of target muscle activity occurred within 8 d of the injection, and paralysis was confirmed using electromyography. No paralysis was seen in the adjacent muscles (abductor profundus) or in positive controls (saline injections). Paralysis occurred more slowly and at lower doses than previously documented for mammals. However, botulinum toxin complex (500 kDa) was used here, whereas previous studies have used purified toxin (150 kDa). Therefore, differences in physiological responses between fish and mammals cannot yet be distinguished from differences caused by the toxin type. However, we note that the toxin complex is less likely to diffuse across muscle fascia (because it is large), which should minimize paralytic effects on adjacent muscles. We suggest that botulinum toxin holds great promise as a chemical denervation agent in functional studies of animal locomotion and feeding behaviors.     

The swimming endurance of threespine sticklebacks, Gasterosteus aculeatus L., from the Afon Rheidol, Wales

The endurance of threespine sticklebacks, Gasterosteus aculeatus , swimming with pectoral fin locomotion at 20° C in a laboratory flume was measured. Each trial lasted a maximum of 480 min. At a speed of 4 body lengths per sec (L s⁻¹) all fish were still swimming at the end of the trial, but endurance decreased at higher speeds. At speeds of 5 or 6 L s⁻¹ (20–30 cm s⁻¹) a few fish still maintained labriform locomotion for the 480 min. However, at a speed of 7 L s⁻¹ all fish furled their pectoral fins and used body and caudal fin propulsion but fatigued rapidly. During sustained swimming, fish could cover distances of 6 km or more. No significant differences between males and females were found.     

Pectoral fin beat frequency predicts oxygen consumption during spontaneous activity in a labriform swimming fish (Embiotoca lateralis)

The objective of this study was to identify kinematic variables correlated with oxygen consumption during spontaneous labriform swimming. Kinematic variables (swimming speed, change of speed, turning angle, turning rate, turning radius and pectoral fin beat frequency) and oxygen consumption (MO₂) of spontaneous swimming in Embiotoca lateralis were measured in a circular arena using video tracking and respirometry, respectively. The main variable influencing MO₂ was pectoral fin beat frequency (r ² = 0.71). No significant relationship was found between swimming speed and pectoral fin beat frequency. Complementary to other methods within biotelemetry such as EMG it is suggested that such correlations of pectoral fin beat frequency may be used to measure the energy requirements of labriform swimming fish such as E. lateralis in the field, but need to be taken with great caution since movement and oxygen consumption patterns are likely to be quite different in field situation compared to a small lab tank. In addition, our methods could be useful to measure metabolic costs of growth and development, or bioassays for possible toxicological effects on fish.     

Beziehungen zwischen Körperbau und Lebensweise bei Blenniidae (Pisces) des Roten Meeres

The muscle and skeleton anatomy of the pectoral, pelvic, and anal fins are described in 3 Salariin Blenniidae: Salarias fasciatus (sublittoral), Istiblennius edentulus (eulittoral), Alticus kirkii (supralittoral). In A. kirkii these organs are adapted to a climbing habit on the steep rocks beyond the water. The results are compared with those found in Periophthalmus.

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Abbildungserklärungen B Basale - Cl Cleithrum - Co Coracoid - Creld Crista cleithri dorsalis - Crele Crista cleithri externa - H Haken an den Lepidotrichen der Ventralia - Lep Lepidotrichen - Pel Postcleithrum - Prsv Processus spinae ventralis - Pt Posttemporale - Rad Radiale - Scl Supracleithrum - SCl Symphyse des Cleithrum - Scp Scapula - abpr M. abductor profundus - adpr M. adductor profundus - arre M. arrector externus - arri M. arrector internus - cord M. coraco-radialis - dep M. depressor (Analis) - dprrd M. depressor radiorum (Pectoralia) - er M. erector - extpr M. extensor proprius - fls M. flexor superficialis - inc M. inclinator - levs M. levator superficialis - mes M. mesoventralis - rtrd M. retractor dorsalis - rtris M. retractor ischii - rtrv M. retractor ventralis Mit Unterstützung der Deutschen Forschungsgemeinschaft.     

Computation of Unsteady Flow Past a Biomimetic Fin

The unsteady hydrodynamics of a biomimetic fin attached to a cylindrical body has been studied numerically using a computational fluid dynamic (CFD) simulator based on an in-house solver of the Navier-Stokes equations, combined with a recently developed multi-block, overset grid method. The fin-body CFD model is based on a mechanical pectoral fin device, which consists of a cylindrical body and an asymmetric fin and can mimic flapping, rowing and feathering motions of the pectoral fins in fishes. First the multi-block, overset grid method incorporated into the NS solver was verified through an extensive study of unsteady flows past a single fin undergoing rowing and feathering motion. Then unsteady flows past the biomimetic fin-body model undergoing the same motions were computed and compared with the measurements of forces of the mechanical pectoral fin, which shows good agreement in both time-varying and time-averaged hydrodynamic forces. The relationship between force generation and vortex dynamics points to the importance of the match in fin kinematics between power and recovery strokes and implies that an optimal selection of parameters of phase lags between and amplitudes of rowing and feathering motions can improve the performance of labriform propulsion in terms of either maximum force generation or minimum mechanical power.     

Locomotion in labrid fishes: implications for habitat use and cross-shelf biogeography on the Great Barrier Reef

Coral reefs exhibit marked zonation patterns within single reefs and across continental shelves. For sessile organisms these zones are often related to wave exposure. We examined the extent to which wave exposure may shape the distribution patterns of fishes. We documented the distribution of 98 species of wrasses and parrotfishes at 33 sites across the Great Barrier Reef. The greatest difference between labrid assemblages was at the habitat level, with exposed reef flats and crests on mid- and outer reefs possessing a distinct faunal assemblage. These exposed sites were dominated by individuals with high pectoral fin aspect ratios, i.e. fishes believed to be capable of lift-based swimming which often achieve high speeds. Overall, there was a strong correlation between estimated swimming performance, as indicated by fin aspect ratio, and degree of water movement. We propose that swimming performance in fishes limits access to high-energy locations and may be a significant factor influencing habitat use and regional biogeography of reef fishes.     

Global ecological success of Thalassoma fishes in extreme coral reef habitats

Phenotypic adaptations can allow organisms to relax abiotic selection and facilitate their ecological success in challenging habitats, yet we have relatively little data for the prevalence of this phenomenon at macroecological scales. Using data on the relative abundance of coral reef wrasses and parrotfishes (f. Labridae) spread across three ocean basins and the Red Sea, we reveal the consistent global dominance of extreme wave‐swept habitats by fishes in the genus Thalassoma, with abundances up to 15 times higher than any other labrid. A key locomotor modification—a winged pectoral fin that facilitates efficient underwater flight in high‐flow environments—is likely to have underpinned this global success, as numerical dominance by Thalassoma was contingent upon the presence of high‐intensity wave energy. The ecological success of the most abundant species also varied with species richness and the presence of congeneric competitors. While several fish taxa have independently evolved winged pectoral fins, Thalassoma appears to have combined efficient high‐speed swimming (to relax abiotic selection) with trophic versatility (to maximize exploitation of rich resources) to exploit and dominate extreme coral reef habitats around the world.     

Locomotor Patterns in the Evolution of Actinopterygian Fishes

SYNOPSIS. Locomotor adaptations in actinopterygian fishes aredescribed for (a) caudal propulsion, used in cruising and sprintswimming, acceleration, and fast turns and (b) median and pairedfin propulsion used for slow swimming and in precise maneuver.Caudal swimming is subdivided into steady (time independent)and unsteady (time dependent acceleration and turning) locomotion. High power caudal propulsion is the major theme in actinopterygianswimming morphology because of its role in predator evasionand food capture. Non-caudal slow swimming appears to be secondaryand is not exploited before the Acanthopterygii. Optimal morphological requirements for unsteady swimming are(a) large caudal fin and general body area, (b) deep caudalpeduncle, often enhanced by posterior dorsal and anal fins,(c) an anterior stabilizing body mass and\or added mass, (d)flexible body and (e) large ratio of muscle mass to body mass.Optimal morphological requirements for steady swimming are (a)high aspect ratio caudal fin, (b) narrow caudal peduncle, (c)small total caudal area, (d) anterior stabilizing body massand added mass, and (e) a stiff body. Small changes in morphologycan have large effects on performance. Exclusive morphological requirements for steady versus unsteadyswimming are partially overcome using collapsible fins, butcompromises remain necessary. Morphologies favoring unsteadyperformance are a recurring theme in actinopterygian evolution.Successive radiations at chondrostean, halecostome and teleosteanlevels are associated with modifications in the axial and caudalskeleton. Strength of ossified structures probably limited maximum propulsionforces early in actinopterygian evolution, so that specializationsfor fast cruising (carangiform and thunmform modes) followedstructural advances especially in the caudal skeleton. No suchlimits apply to eel-like forms which consequently recur in successiveactinopterygian radiations. Slow swimming using mainly non-caudal propulsion probably firstoccurred among neopterygians in association with reduced andneutral buoyancy. Slow swimming adaptations can add to and extendthe scope of caudal swimming, but specialization is associatedwith reduced caudal swimming performance. Marked exploitationof slow swimming opportunities does not occur prior to the anterodorsallocation of pectoral and pelvic girdles and the vertical rotationof the base of the pectoral fin, as found in the Acanthopterygii.     

The evolution of underwater flight: The redistribution of pectoral fin rays,in manta rays and their relatives (Myliobatidae)

Batoids are a diverse clade of flat cartilaginous fishes that occur primarily in benthic marine habitats. The skates and rays typically use their flexible pectoral fins for feeding and propulsion via undulatory swimming. However, two groups of rays have adopted a pelagic or bentho‐pelagic lifestyle and utilize oscillatory swimming—the Myliobatidae and Gymnuridae. The myliobatids have evolved cephalic lobes, anteriorly extended appendages that are optimized for feeding, while their pectoral fins exhibit several modifications that likely arose in association with functional optimization of pelagic cruising via oscillatory flight. Here, we examine variation in fin ray distribution and ontogenetic timing of fin ray development in batoid pectoral fins in an evolutionary context using the following methods: radiography, computed tomography, dissections, and cleared and stained specimens. We propose an index for characterizing variation in the distribution of pectoral fin rays. While undulatory swimmers exhibit symmetry or slight anterior bias, we found a posterior shift in the distribution of fin rays that arose in two distinct lineages in association with oscillatory swimming. Undulatory and oscillatory swimmers occupy nonoverlapping morphospace with respect to fin ray distribution illustrating significant remodeling of pectoral fins in oscillatory swimmers. Further, we describe a derived skeletal feature in anterior pectoral fins of the Myliobatidae that is likely associated with optimization of oscillatory swimming. By examining the distribution of fin rays with clearly defined articulation points, we were able to infer evolutionary trends and body plan remodeling associated with invasion of the pelagic environment. Finally, we found that the number and distribution of fin rays is set early in development in the little skate, round stingray, and cownose ray, suggesting that fin ray counts from specimens after birth or hatching are representative of adults and therefore comparable among species.     

Design and Implementation of Paired Pectoral Fins Locomotion of Labriform Fish Applied to a Fish Robot

In present,there are increasing interests in the research on mechanical and control system of underwater vehicles.Theseongoing research efforts are motivated by more pervasive applications of such vehicles including seabed oil and gas explorations,scientific deep ocean surveys,military purposes,ecological and water environmental studies,and also entertainments.However,the performance of underwater vehicles with screw type propellers is not prospective in terms of its efficiency andmaneuverability.The main weaknesses of this kind of propellers are the production of vortices and sudden generation of thrustforces which make the control of the position and motion difficult.On the other hand,fishes and other aquatic animals are efficient swimmers,posses high maneuverability,are able to followtrajectories,can efficiently stabilize themselves in currents and surges,create less wakes than currently used underwater vehicle,and also have a noiseless propulsion.The fish’s locomotion mechanism is mainly controlled by its caudal fin and paired pectoralfins.They are classified into Body and/or Caudal Fin(BCF)and Median and/or paired Pectoral Fins(MPF).The study of highlyefficient swimming mechanisms of fish can inspire a better underwater vehicles thruster design and its mechanism.There are few studies on underwater vehicles or fish robots using paired pectoral fins as thruster.The work presented in thispaper represents a contribution in this area covering study,design and implementation of locomotion mechanisms of pairedpectoral fins in a fish robot.The performance and viability of the biomimetic method for underwater vehicles are highlightedthrough in-water experiment of a robotic fish.     

Morphological and histochemical characterization of the pectoral fin muscle of batoids

Batoids differ from other elasmobranch fishes in that they possess dorsoventrally flattened bodies with enlarged muscled pectoral fins. Most batoids also swim using either of two modes of locomotion: undulation or oscillation of the pectoral fins. In other elasmobranchs (e.g., sharks), the main locomotory muscle is located in the axial myotome; in contrast, the main locomotory muscle in batoids is found in the enlarged pectoral fins. The pectoral fin muscles of sharks have a simple structure, confined to the base of the fin; however, little to no data are available on the more complex musculature within the pectoral fins of batoids. Understanding the types of fibers and their arrangement within the pectoral fins may elucidate how batoid fishes are able to utilize such unique swimming modes. In the present study, histochemical methods including succinate dehydrogenase (SDH) and immunofluoresence were used to determine the different fiber types comprising these muscles in three batoid species: Atlantic stingray (Dasyatis sabina), ocellate river stingray (Potamotrygon motoro) and cownose ray (Rhinoptera bonasus). All three species had muscles comprised of two muscle fiber types (slow-red and fast-white). The undulatory species, D. sabina and P. motoro, had a larger proportion of fast-white muscle fibers compared to the oscillatory species, R. bonasus. The muscle fiber sizes were similar between each species, though generally smaller compared to the axial musculature in other elasmobranch fishes. These results suggest that batoid locomotion can be distinguished using muscle fiber type proportions. Undulatory species are more benthic with fast-white fibers allowing them to contract their muscles quickly, as a possible means of escape from potential predators. Oscillatory species are pelagic and are known to migrate long distances with muscles using slow-red fibers to aid in sustained swimming.     

The brain organization of butterflyfishes

Synopsis The encephalization indices of angelfishes (Pomacanthidae) and butterflyfishes (Chaetodontidae) are typical of advanced perciform fishes: both families lie in the upper part of the polygon of teleost indices. The chaetodontids seem to be a little more encephalized than pomacanthids. The general morphology of the brains in both families is very similar: small olfactory bulbs, large optic tectum and a cerebellum which covers the brain structures in front of it like a cap. This morphology is shared by another family of the coral reef biotope, the Acanthuridae. The histological architecture is also typical of advanced teleosts, with a cortex-like pallium, a laminated nucleus geniculatus (= pretectalis superficialis), a complex valvula cerebelli and a corpus glomerulosum with a clear neuropile centre. The quantitative analysis of the main subdivisions of the brain, either from relative volumes or from indices, shows small olfactory bulbs (microsmy) but important telencephalic and diencephalic centres, large tectal centres (vision) and large cerebellum (precise locomotion). Many of these peculiarities are shared by other fishes inhabiting coral reefs. The differences between the two families seem to be primarily correlated with food habits: the angelfishes, which are sponge-feeders and may have an overweight due to the ballast of the sponge-skeleton in their digestive tract, and which do not need either such good vision or such precise locomotion to pick up their prey, could be a little less encephalized than the butterflyfishes.     

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

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