Distribution, swimming physiology, and swimming mechanics of brief squid Lolliguncula brevis

Squids are thought to have physiological and locomotive deficiencies that put them at a competitive disadvantage to fishes and exclude them from inshore, highly variable environments that are rich in nektonic fauna. However, brief squid Lolliguncula brevis may be a notable exception. Trawl surveys revealed that L. brevis, particularly juveniles \u3c6 cm dorsal mantle length (DML), are abundant in the Chesapeake Bay, especially when salinity and water temperature are high, and tolerate a wide range of physical conditions relative to other cephalopods. L. brevis is also different from other cephalopods examined previously because its pattern of oxygen consumption as a function of velocity was found to be parabolic and thus similar to aerial flight, and its swimming costs were competitive with ecologically equivalent fishes. Power-speed curves derived from video footage of swimming squid and hydrodynamic force calculations also were parabolic in shape, with high costs both at low and high speeds because of power requirements for lift generation and overcoming drag, respectively. L. brevis employed various behaviors to increase swimming efficiency and compensate for negative buoyancy, such as swimming in various orientations (e.g., arms-first and tail-first), altering angles of attack of the mantle, arms, and funnel, and using fin activity. Fin motion, which could not be characterized exclusively as drag- or lift-based propulsion, was used over 50--95% of the sustained speed range and provided as much as 78% of the vertical and 55% of the horizontal thrust. Small squid (\u3c3.0 cm DML) used different swimming strategies than larger squid possibly to maximize the benefits of toroidal induction, and aerobic efficiency curves indicated that squid 3--5 cm. DML are most efficient. Brief squid also may take advantage of unsteady phenomena, such as attached vortices, for added lift and thrust. Furthermore, an electromyographic study revealed that L. brevis uses different circular muscle layers for various speeds and like fish has muscular gears , suggesting that there is specialization and efficient use of locomotive muscle in some cephalopods. Therefore, the presumption that squids are inescapably constrained by a second-rate propulsive system and physiological deficiencies is not applicable to L. brevis

Small-Scale Patterns of Recruitment On A Constructed Intertidal Reef: The Role of Spatial Refugia

Traditional oyster repletion activities have utilized a two-dimensional approach to shell (substrate) deployment to attain maximal coverage in subtidal locations with little consideration for optimal thickness of deployed shell and tidal elevation. Vertical dimensionality may play a vital role, however, in the establishment and persistence of oyster communities. Therefore, a three-dimensional oyster reef was constructed in the Piankatank River, Virginia, and settlement and mortality patterns of oysters were recorded from June of 1993 through September of 1994.https://scholarworks.wm.edu/vimsbooks/1087/thumbnail.jp

Ontogenetic Investigation of Underwater Hearing Capabilities in Loggerhead Sea Turtles (Caretta caretta) Using a Dual Testing Approach

Sea turtles reside in different acoustic environments with each life history stage and may have different hearing capacity throughout ontogeny. For this study, two independent yet complementary techniques for hearing assessment, i.e. behavioral and electrophysiological audiometry, were employed to (1) measure hearing in post-hatchling and juvenile loggerhead sea turtles Caretta caretta (19-62 cm straight carapace length) to determine whether these migratory turtles exhibit an ontogenetic shift in underwater auditory detection and (2) evaluate whether hearing frequency range and threshold sensitivity are consistent in behavioral and electrophysiological tests. Behavioral trials first required training turtles to respond to known frequencies, a multi-stage, time-intensive process, and then recording their behavior when they were presented with sound stimuli from an underwater speaker using a two-response forced-choice paradigm. Electrophysiological experiments involved submerging restrained, fully conscious turtles just below the air-water interface and recording auditory evoked potentials (AEPs) when sound stimuli were presented using an underwater speaker. No significant differences in behavior-derived auditory thresholds or AEP-derived auditory thresholds were detected between post-hatchling and juvenile sea turtles. While hearing frequency range (50-1000/1100 Hz) and highest sensitivity (100-400 Hz) were consistent in audiograms pooled by size class for both behavior and AEP experiments, both post-hatchlings and juveniles had significantly higher AEP-derived than behavior-derived auditory thresholds, indicating that behavioral assessment is a more sensitive testing approach. The results from this study suggest that post-hatchling and juvenile loggerhead sea turtles are low-frequency specialists, exhibiting little differences in threshold sensitivity and frequency bandwidth despite residence in acoustically distinct environments throughout ontogeny

Lateral Line Analogue Aids Vision in Successful Predator Evasion for the Brief Squid, Lolliguncula Brevis

Cephalopods have visual and mechanoreception systems that may be employed to sense and respond to an approaching predator. While vision presumably plays the dominant role, the importance of the lateral line analogue for predator evasion has not been examined in cephalopods. To test the respective roles of vision and the lateral line analogue, brief squid, Lolliguncula brevis, were observed in the presence of summer flounder, Paralichthys dentatus, under light and dark conditions with their lateral line analogue intact and ablated. Hair cell ablation was achieved through a pharmacological technique used for the first time on a cephalopod. The proportion of predator-prey interactions survived was significantly higher in the light non-ablated and light ablated groups compared with the dark ablated group. The mean number of interactions survived varied across treatment groups with the light non-ablated group having significantly more success than the light ablated, dark non-ablated and dark ablated groups. These findings demonstrate that although vision is the primary sense, the lateral line analogue also contributes to predator evasion in squid

Hydrodynamic Fin Function of Brief Squid, Lolliguncula Brevis

Although the pulsed jet is often considered the foundation of a squid\u27s locomotive system, the lateral fins also probably play an important role in swimming, potentially providing thrust, lift and dynamic stability as needed. Fin morphology and movement vary greatly among squid species, but the locomotive role of the fins is not well understood. To begin to elucidate the locomotive role of the fins in squids, fin hydrodynamics were studied in the brief squid Lolliguncula brevis, a species that exhibits a wide range of fin movements depending on swimming speed. Individual squid were trained to swim in both the arms-first and tail-first orientations against currents in a water tunnel seeded with light-reflective particles. Particle-laden water around the fins was illuminated with lasers and videotaped so that flow dynamics around the fins could be analyzed using digital particle image velocimetry (DPIV). Time-averaged forces generated by the fin were quantified from vorticity fields of the fin wake. During the low swimming speeds considered in this study [\u3c2.5 dorsal mantle lengths (DML) per second], L. brevis exhibited four unique fin wake patterns, each with distinctive vortical structures: (1) fin mode I, in which one vortex is shed with each downstroke, generally occurring at low speeds; (2) fin mode II, an undulatory mode in which a continuous linked chain of vortices is produced; (3) fin mode III, in which one vortex is shed with each downstroke and upstroke, and; (4) fin mode IV, in which a discontinuous chain of linked double vortex structures is produced. All modes were detected during tail-first swimming but only fin modes II and III were observed during arms-first swimming. The fins produced horizontal and vertical forces of varying degrees depending on stroke phase, swimming speed, and swimming orientation. During tail-first swimming, the fins functioned primarily as stabilizers at low speeds before shifting to propulsors as speed increased, all while generating net lift. During arms-first swimming, the fins primarily provided lift with thrust production playing a reduced role. These results demonstrate the lateral fins are an integral component of the complex locomotive system of L. brevis, producing lift and thrust forces through different locomotive modes

Swimming mechanics and behavior of the shallow-water brief squid Lolliguncula brevis

Although squid are among the most versatile swimmers and rely on a unique locomotor system, little is known about the swimming mechanics and behavior of most squid, especially those that swim at low speeds in inshore waters. Shallow-water brief squid Lolliguncula brevis, ranging in size from 1.8 to 8.9 cm in dorsal mantle length (DML), were placed in flumes and videotaped, and the data were analyzed using motion-analysis equipment. Flow visualization and force measurement experiments were also performed in water tunnels. Mean critical swimming speeds (Ucrit) ranged from 15.3 to 22.8 cm s–1, and mean transition speeds (Ut; the speed above which squid swim exclusively in a tail-first orientation) varied from 9.0 to 15.3 cm s–1. At low speeds, negatively buoyant brief squid generated lift and/or improved stability by positioning the mantle and arms at high angles of attack, directing high-speed jets downwards (angles \u3e50°) and using fin activity. To reduce drag at high speeds, the squid decreased angles of attack and swam tail-first. Fin motion, which could not be characterized exclusively as drag- or lift-based propulsion, was used over 50–95 % of the sustained speed range and provided as much as 83.8 % of the vertical and 55.1 % of the horizontal thrust. Small squid (DML) used different swimming strategies from those of larger squid, possibly to maximize thrust benefits from vortex ring formation. Furthermore, brief squid employed various unsteady behaviors, such as manipulating funnel diameter during jetting, altering arm position and swimming in different orientations, to boost swimming performance. These results demonstrate that locomotion in slow-swimming squid is complex, involving intricate spatial and temporal interactions between the mantle, fins, arms and funnel

Hydrodynamic stability of swimming in ostraciid fishes: role of the carapace in the smooth trunkfish Lactophrys triqueter (Teleostei: Ostraciidae)

The hydrodynamic bases for the stability of locomotory motions in fishes are poorly understood, even for those fishes, such as the rigid-bodied smooth trunkfish Lactophrys triqueter, that exhibit unusually small amplitude recoil movements during rectilinear swimming. We have studied the role played by the bony carapace of the smooth trunkfish in generating trimming forces that self-correct for instabilities. The flow patterns, forces and moments on and around anatomically exact, smooth trunkfish models positioned at both pitching and yawing angles of attack were investigated using three methods: digital particle image velocimetry (DPIV), pressure distribution measurements, and force balance measurements. Models positioned at various pitching angles of attack within a flow tunnel produced well-developed counter-rotating vortices along the ventro-lateral keels. The vortices developed first at the anterior edges of the ventro-lateral keels, grew posteriorly along the carapace, and reached maximum circulation at the posterior edge of the carapace. The vortical flow increased in strength as pitching angles of attack deviated from 0°, and was located above the keels at positive angles of attack and below them at negative angles of attack. Variation of yawing angles of attack resulted in prominent dorsal and ventral vortices developing at far-field locations of the carapace; far-field vortices intensified posteriorly and as angles of attack deviated from 0°. Pressure distribution results were consistent with the DPIV findings, with areas of low pressure correlating well with regions of attached, concentrated vorticity. Lift coefficients of boxfish models were similar to lift coefficients of delta wings, devices that also generate lift through vortex generation. Furthermore, nose-down and nose-up pitching moments about the center of mass were detected at positive and negative pitching angles of attack, respectively. The three complementary experimental approaches all indicate that the carapace of the smooth trunkfish effectively generates self-correcting forces for pitching and yawing motions — a characteristic that is advantageous for the highly variable velocity fields experienced by trunkfish in their complex aquatic environment. All important morphological features of the carapace contribute to producing the hydrodynamic stability of swimming trajectories in this species

Turning Performance in Squid and Cuttlefish: Unique Dual-Mode, Muscular Hydrostatic Systems

Although steady swimming has received considerable attention in prior studies, unsteady swimming movements represent a larger portion of many aquatic animals\u27 locomotive repertoire and have not been examined extensively. Squids and cuttlefishes are cephalopods with unique muscular hydrostat-driven, dual-mode propulsive systems involving paired fins and a pulsed jet. These animals exhibit a wide range of swimming behavior, but turning performance has not been examined quantitatively. Brief squid, Lolliguncula brevis, and dwarf cuttlefish, Sepia bandensis, were filmed during turns using high-speed cameras. Kinematic features were tracked, including the length-specific radius of the turn (R/L), a measure of maneuverability, and angular velocity (ω), a measure of agility. Both L. brevis and S. bandensis demonstrated high maneuverability, with (R/L)min values of 3.4x10(-3)+/- 5.9x10(-4) and 1.2x10(-3)+/- 4.7x10(-4) (mean +/- s.e.m.), respectively, which are the lowest measures of R/L reported for any aquatic taxa. Lolliguncula brevis exhibited higher agility than S. bandensis (ωa,max=725.8 versus 485.0 deg s-1), and both cephalopods have intermediate agility when compared with flexible-bodied and rigid-bodied nekton of similar size, reflecting their hybrid body architecture. In L. brevis, jet flows were the principal driver of angular velocity. Asymmetric fin motions played a reduced role, and arm wrapping increased turning performance to varying degrees depending on the species. This study indicates that coordination between the jet and fins is important for turning performance, with L. brevis achieving faster turns than S. bandensis and S. bandensis achieving tighter, more controlled turns than L. brevis

Aerobic respiratory costs of swimming in the negatively buoyant brief squid Lolliguncula brevis

Because of the inherent inefficiency of jet propulsion, squid are considered to be at a competitive disadvantage compared with fishes, which generally depend on forms of undulatory/oscillatory locomotion. Some squid, such as the brief squid Lolliguncula brevis, swim at low speeds in shallow-water complex environments, relying heavily on fin activity. Consequently, their swimming costs may be lower than those of the faster, more pelagic squid studied previously and competitive with those of ecologically relevant fishes. To examine aerobic respiratory swimming costs, O2 consumption rates were measured for L. brevis of various sizes (2–9 cm dorsal mantle length, DML) swimming over a range of speeds (3–30 cm s–1) in swim tunnel respirometers, while their behavior was videotaped. Using kinematic data from swimming squid and force data from models, power curves were also generated. Many squid demonstrated partial (J-shaped) or full (U-shaped) parabolic patterns of O2 consumption rate as a function of swimming speed, with O2 consumption minima at 0.5–1.5 DML s–1. Power curves derived from hydrodynamic data plotted as a function of swimming speed were also parabolic, with power minima at 1.2–1.7 DML s–1. The parabolic relationship between O2 consumption rate/power and speed, which is also found in aerial flyers such as birds, bats and insects but rarely in aquatic swimmers because of the difficulties associated with low-speed respirometry, is the result of the high cost of generating lift and maintaining stability at low speeds and overcoming drag at high speeds. L. brevis has a lower rate of O2 consumption than the squid Illex illecebrosus and Loligo opalescens studied in swim tunnel respirometers and is energetically competitive (especially at O2 consumption minima) with fishes, such as striped bass, mullet and flounder. Therefore, the results of this study indicate that, like aerial flyers, some negatively buoyant nekton have parabolic patterns of O2 consumption rate/power as a function of speed and that certain shallow-water squid using considerable fin activity have swimming costs that are competitive with those of ecologically relevant fishes