Propulsion in Cubomedusae: Mechanisms and Utility

Evolutionary constraints which limit the forces produced during bell contractions of medusae affect the overall medusan morphospace such that jet propulsion is limited to only small medusae. Cubomedusae, which often possess large prolate bells and are thought to swim via jet propulsion, appear to violate the theoretical constraints which determine the medusan morphospace. To examine propulsion by cubomedusae, we quantified size related changes in wake dynamics, bell shape, swimming and turning kinematics of two species of cubomedusae, Chironex fleckeri and Chiropsella bronzie. During growth, these cubomedusae transitioned from using jet propulsion at smaller sizes to a rowing-jetting hybrid mode of propulsion at larger sizes. Simple modifications in the flexibility and kinematics of their velarium appeared to be sufficient to alter their propulsive mode. Turning occurs during both bell contraction and expansion and is achieved by generating asymmetric vortex structures during both stages of the swimming cycle. Swimming characteristics were considered in conjunction with the unique foraging strategy used by cubomedusae.     

Ontogenetic changes in the bell morphology and kinematics and swimming behavior of rowing medusae: the special case of the limnomedusa Liriope tetraphylla

Swimming animals may experience significant changes in the Reynolds number (Re) of their surrounding fluid flows throughout ontogeny. Many medusae experience Re environments with significant viscous forces as small juveniles but inertially dominated Re environments as adults. These different environments may affect their propulsive strategies. In particular, rowing, a propulsive strategy with ecological advantages for large adults, may be constrained by viscosity for small juvenile medusae. We examined changes in the bell morphology and swimming kinematics of the limnomedusa Liriope tetraphylla at different stages of development. L. tetraphylla maintained an oblate bell (fineness ratio ≈ 0.5-0.6), large velar aperture ratio (R(v) ≈ 0.5-0.8), and rapid bell kinematics throughout development. These traits enabled it to use rowing propulsion at all stages except the very smallest sizes observed (diameter = 0.14 cm). During the juvenile stage, very rapid bell kinematics served to increase Re sufficiently for rowing propulsion. Other taxa that use rowing propulsion as adults, such as leptomedusae and scyphomedusae, typically utilize different propulsive strategies as small juveniles to function in low Re environments. We compared the performance values of the different propulsive modes observed among juvenile medusae.     

Transitions in morphology, nematocyst distribution, fluid motions, and prey capture during development of the scyphomedusa Cyanea capillata

Like that of most scyphozoans, the ontogeny of Cyanea capillata medusae involves substantive alterations in feeding structures and mechanics. We used video and optical microscopy approaches to quantify these ontogenetic changes in morphology, flow, and feeding of C. capillata medusae. We found that alterations in gross morphology and nematocyst distributions coincided with a shift from prey capture on the manubrium or lappets of ephyrae (bell diameter 0.2-0.4 cm) to capture primarily on the tentacles in adult medusae (diameter >1.0 cm). These changes occurred within a hydrodynamic framework that itself changed due to medusan growth. Viscous forces were important in flows around small ephyrae (maximum Re <10(1)), whereas viscosity was less influential in the inertially dominated flows around adult medusae (Re > 10(2)). The relative timing of these events indicates that ontogenetic processes are closely synchronized with alterations in the hydrodynamic environment within which C. capillata medusae develop.     

Reynolds number limits for jet propulsion: a numerical study of simplified jellyfish

The Scallop theorem states that reciprocal methods of locomotion, such as jet propulsion or paddling, will not work in Stokes flow (Reynolds number=0). In nature the effective limit of jet propulsion is still in the range where inertial forces are significant. It appears that almost all animals that use jet propulsion swim at Reynolds numbers (Re) of about 5 or more. Juvenile squid and octopods hatch from the egg already swimming in this inertial regime. Juvenile jellyfish, or ephyrae, break off from polyps swimming at Re greater than 5. Many other organisms, such as scallops, rarely swim at Re less than 100. The limitations of jet propulsion at intermediate Re is explored here using the immersed boundary method to solve the 2D Navier-Stokes equations coupled to the motion of a simplified jellyfish. The contraction and expansion kinematics are prescribed, but the forward and backward swimming motions of the idealized jellyfish are emergent properties determined by the resulting fluid dynamics. Simulations are performed for both an oblate bell shape using a paddling mode of swimming and a prolate bell shape using jet propulsion. Average forward velocities and work put into the system are calculated for Re between 1 and 320. The results show that forward velocities rapidly decay with decreasing Re for all bell shapes when Re<10. Similarly, the work required to generate the pulsing motion increases significantly for Re<10. When compared to actual organisms, the swimming velocities and vortex separation patterns for the model prolate agree with those observed in Nemopsis bachei. The forward swimming velocities of the model oblate jellyfish after two pulse cycles are comparable to those reported for Aurelia aurita, but discrepancies are observed in the vortex dynamics between when the 2D model oblate jellyfish and the organism. This discrepancy is likely due to a combination of the differences between the 3D reality of the jellyfish and the 2D simplification, as well as the rigidity of the time varying geometry imposed by the idealized model.     

Swimming dynamics and propulsive efficiency of squids throughout ontogeny

Squids encounter vastly different flow regimes throughout ontogeny as they undergo critical morphological changes to their two locomotive systems: the fins and jet. Squid hatchlings (paralarvae) operate at low and intermediate Reynolds numbers (Re) and typically have rounded bodies, small fins, and relatively large funnel apertures, whereas juveniles and adults operate at higher Re and generally have more streamlined bodies, larger fins, and relatively small funnel apertures. These morphological changes and varying flow conditions affect swimming performance in squids. To determine how swimming dynamics and propulsive efficiency change throughout ontogeny, digital particle image velocimetry (DPIV) and kinematic data were collected from an ontogenetic range of long-finned squid Doryteuthis pealeii and brief squid Lolliguncula brevis swimming in a holding chamber or water tunnel (Re = 20-20 000). Jet and fin wake bulk properties were quantified, and propulsive efficiency was computed based on measurements of impulse and excess kinetic energy in the wakes. Paralarvae relied predominantly on a vertically directed, high frequency, low velocity jet as they bobbed up and down in the water column. Although some spherical vortex rings were observed, most paralarval jets consisted of an elongated vortical region of variable length with no clear pinch-off of a vortex ring from the trailing tail component. Compared with paralarvae, juvenile and adult squid exhibited a more diverse range of swimming strategies, involving greater overall locomotive fin reliance and multiple fin and jet wake modes with better defined vortex rings. Despite greater locomotive flexibility, jet propulsive efficiency of juveniles/adults was significantly lower than that of paralarvae, even when juvenile/adults employed their highest efficiency jet mode involving the production of periodic isolated vortex rings with each jet pulse. When the fins were considered together with the jet for several juvenile/adult swimming sequences, overall propulsive efficiency increased, suggesting that fin contributions are important and should not be overlooked in analyses of the swimming performance of squids. The fins produced significant thrust and consistently had higher propulsive efficiency than did the jet. One particularly important area of future study is the determination of coordinated jet/fin wake modes that have the greatest impact on propulsive efficiency. Although such research would be technically challenging, requiring new, powerful, 3D approaches, it is necessary for a more comprehensive assessment of propulsive efficiency of the squid dual-mode locomotive system.     

In situ time budgets of the scyphomedusae Aurelia aurita, Cyanea sp., and Chrysaora quinquecirrha

The in situ behavior of three scyphomedusan species was videorecorded by scuba divers in natural daytime lighting with minimalinterference to the medusae. The mean percentage of time thatindividual medusae spent swimming ranged from 93 to 100%. Therewere no significant differences in the percent time spent swimmingbetween life stages of a species (ephyra, adult) or betweenspecies. The predominance of swimming activity by medusae indicatesthat swimming, and hence the creation of fluid motions responsiblefor prey entrainment and capture, plays a widespread functionalrole in feeding by scyphomedusae.     

Size-based Prey Selectivity and Dietary Shifts in the Jellyfish, Aurelia aurita

Gut content analyses on field-caught Aurelia aurita showed bothquantitative and qualitative change in diet as a function ofmedusa size. Larger medusae tended towards greater numbersand diversity of prey (up to 1550 individual prey representingas many as 13 different prey groups). We also found that medusasize was a good predictor of prey diversity recovered from themedusa gut. While a shift toward greater prey diversity inlarger medusae might be explained by increased contact rateswith 'rare' prey taxa, we also found size-based prey selectivitychanges in A. aurita. We used in situ gut content data to describeselectivity by A. aurita for three prey types representing varyingdegrees of swimming or escape velocity. Fish eggs were usedas a non-swiming prey, and small (     

Foraging mode affects the evolution of egg size in generalist predators embedded in complex food webs

Ecological networks incorporate myriad biotic interactions that determine the selection pressures experienced by the embedded populations. We argue that within food webs, the negative scaling of abundance with body mass and foraging theory predict that the selective advantages of larger egg size should be smaller for sit‐and‐wait than active‐hunting generalist predators, leading to the evolution of a difference in egg size between them. Because body mass usually scales negatively with predator abundance and constrains predation rate, slightly increasing egg mass should simultaneously allow offspring to feed on more prey and escape from more predators. However, the benefits of larger offspring would be relatively smaller for sit‐and‐wait predators because (i) due to their lower mobility, encounters with other predators are less common, and (ii) they usually employ a set of alternative hunting strategies that help to subdue relatively larger prey. On the other hand, for active predators, which need to confront prey as they find them, body‐size differences may be more important in subduing prey. This difference in benefits should lead to the evolution of larger egg sizes in active‐hunting relative to sit‐and‐wait predators. This prediction was confirmed by a phylogenetically controlled analysis of 268 spider species, supporting the view that the structure of ecological networks may serve to predict relevant selective pressures acting on key life history traits.     

New trace fossil evidence for eurypterid swimming behaviour

We describe a recently discovered trace fossil from a eurypterid Konservat‐Lagerstätte in the upper Silurian Tonoloway Formation of Pennsylvania, and formally describe contemporaneous traces from the Williamsville Formation Lagerstätte of Ontario. The traces from both localities are assigned here to Arcuites bertiensis igen. et isp. nov. Based on comparisons with previously described eurypterid trackways, neoichnological experiments, and the co‐occurrence with eurypterid remains, Arcuites is interpreted as having been made by the swimming leg (sixth prosomal appendage) of swimming juvenile to adult eurypteroid eurypterids, and represents the first unambiguous trace fossil evidence for eurypterid swimming behaviour. The morphology of Arcuites indicates that eurypteroid eurypterids swam using drag‐based rowing, whereby the animal propelled itself forward by moving its oar blade‐like swimming paddles in an in‐phase backstroke. Arcuites morphology also indicates that the eurypteroid swimming appendage had a greater degree of movement than was previously suggested, and a revised rowing model is proposed. Differences in the abundance of A. bertiensis in the Tonoloway and Williamsville formations suggest a bathymetric control on eurypterid swimming behaviour and trace production. The association of Arcuites with eurypterid body fossils in both units indicates that these Lagerstätten were autochthonous assemblages and provides additional evidence for eurypterid inhabitation of shallow subtidal marine environments in the late Silurian.     

Foreflipper propulsion in the California sea lion, Zalophus californianus

The family Otariidae comprises the only group of marine mammals that habitually use their pectoral appendages to generate propulsive forces during swimming. This method of propulsion was examined in the California sea lion ( Zalophus californianus ), a representative member of the family. High-speed films were taken as a sea lion swam against a water current generated inside a large flow channel. Thrust production was determined by examining the body's movement at various stages of the propulsive cycle. Sea lions generate thrust continuously throughout the stroke. Over its initial three-quarters, foreflippers act as hydrofoils creating forward thrust and lift as they move vertically through the water. Thrust production is greatest, however, near the end of the stroke, when flippers are used as paddles and are oriented broad side to the oncoming flow. The force generated by this three-phased system of propulsion is likely to be greater than that attainable by either an exclusively lift-based hydrofoil or drag-based paddling style of swimming.
The kinematic changes that enable sea lions to change speed were also investigated. Film records revealed that stroke amplitude became greater with speed, although total stroke duration remained essentially constant. Sea lions increase stroke frequency with velocity but large variations in the measured values suggest that changes in amplitude and flipper angle of attack are also important parameters for modulating swimming speed.     

Phylogeny of Medusozoa and the evolution of cnidarian life cycles

To investigate the evolution of cnidarian life cycles, data from the small subunit of the ribosome are used to derive a phylogenetic hypothesis for Medusozoa. These data indicate that Cnidaria is monophyletic and composed of Anthozoa and Medusozoa. While Cubozoa and Hydrozoa are well supported clades, Scyphozoa appears to be paraphyletic. Stauromedusae is possibly the sister group of either Cubozoa or all other medusozoans. The phylogenetic results suggest that: the polyp probably preceded the medusa in the evolution of Cnidaria; within Hydrozoa, medusa development involving the entocodon is ancestral; within Trachylina, the polyp was lost and subsequently regained in the parasitic narcomedusans; within Siphonophorae, the float originated prior to swimming bells; stauromedusans are not likely to be descended from ancestors that produced medusae by strobilation; and cubozoan polyps are simplified from those of their ancestors, which possessed polyps with gastric septa and four mesogleal muscle bands and peristomial pits.     

Motility of zooplankton: fitness, foraging and predation

The relative fitness of planktonic organisms foraging underthe risk of predation is examined in terms of their swimmingspeed, path geometry and jump frequency. Fitness is quantifiedin terms of encounter and ingestion of prey, respiration andenergy cost associated with swimming and mortality due to encounterswith predators. It is shown that a convoluted swimming pathin the form of meanders, zigzags or spirals confers greaterfitness than swimming along a straight path. Optimal path configurationis such that the length-scale of the path-meanders is commensuratewith an organism's detection radius to prey, which in turn scaleswith the size of the organism. Optimal swimming speed for acruise-feeding organism decreases with increasing prey concentrationand increasing risk due to ambush predators. For ambush feedingon motile prey, a benefit is gained by periodically moving toa new location. The time spent swimming is largely a functionof energetic costs, whereas the time spent feeding is stronglycontrolled by prey concentration and the risk posed, in turn,by ambush predators. These predictions are supported by observationsdrawn from the literature.     

Effect of body roll amplitude and arm rotation speed on propulsion of arm amputee swimmers

Only a limited amount of research has gone into evaluating the contribution made by the upper arm to the propulsion of elite swimmers with an amputation at elbow level. With assistance of computational fluid dynamics (CFD) modelling, the swimming technique of competitive arm amputee swimmers can be assessed through numerical simulations which test the effect of various parameters on the effectiveness of the swimming propulsion.This numerical study investigates the effect of body roll amplitude and of upper arm rotation speed on the propulsion of an arm amputee swimmer, at different mean swimming speeds. Various test cases are simulated resulting in a thorough analysis of the complex body/fluid interaction with a detailed quantitative assessment of the effect of the variation of each parameter on the arm propulsion. It is found that a body roll movement with an amplitude of 45° enhances greatly the propulsive contribution from the upper arm with an increase of about 70% in the propulsive force compared to the no roll condition. An increase in the angular velocity of the upper arm also leads to a concomitant increase in the propulsive forces produced by the arm.Such results have direct implications for competitive arm amputee front crawl swimmers and for those who coach them. One important message that emerges in this present work is that there exists, for any given swimming speed, a minimum angular velocity at which the upper arm must be rotated to generate effective propulsion. Below this velocity, the upper arm will experience a net resistive drag force which adversely affects swimming performance.     

Morphological adaptations to marine life in snakes

We investigated morphological adaptations to aquatic life within animals that exhibit a structurally simple, elongate body form, i.e., snakes. This linear body plan should impose different biomechanical constraints than the classical streamlined body shape associated with propulsion by fins, feet, or wings. Our measurements of general body shape of terrestrial, amphibious, and marine snakes (all from the same phylogenetic lineage, the Elapidae) show that seasnakes display specialized morphological attributes for life in water. Most notably, the cross‐sectional body shape is circular in terrestrial snakes but dorso‐ventrally elongated in seasnakes (due to a prominent ventral keel); amphibious species (sea kraits) exhibit an intermediate shape. The tail of amphibious and marine species (a major propulsive structure during swimming) is higher and thinner than in terrestrial snakes (i.e., paddle‐shaped) but shorter relative to body length. The evolution of a laterally compressed shape has been achieved by an increase in body height rather than a decrease in body width, possibly reflecting selection for more effective propulsive thrust, and for an ability to maintain hydrodynamic efficiency despite the minor bodily distension inevitably caused by prey items and developing offspring. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc     

Field observations of four Aurelia labiata jellyfish behaviours: swimming down in response to low salinity pre-empted swimming up in response to touch,but animal and plant materials were captured equally

Jellyfish live in complex environments and must continually make behavioural choices. In field observations, adult Aurelia labiata were confronted with a conflict between swimming up elicited by touch of the manubrium and swimming down elicited by low salinity. Following a touch, downward-swimming medusae (1.5–2.0 m deep) turned and swam to within 0.5 m of the surface when the salinity in the top 1.5 m of the water column was greater than 20 ppt but medusae uniformly refused to swim up into the top 1.25 m when the salinity was less than 20 ppt even after being touched three times. The central nervous system of A. labiata appears to have neural circuitry that specifies their response when medusae encounter stimuli that elicit incompatible behaviours. Upward-swimming adult medusae had animal, vegetable or cellulose (paper) material dispersed ahead of them. Medusae captured each material on the bell margin and transported it to a gastric pouch. Medusae displayed only minor behavioural differences in the process. Having sensory, neural and muscular systems organized to capture and pass to the stomach, a huge variety of materials allows medusae to survive in different seasons and environments.     

The long‐term impacts of predators on prey: inducible defenses,population dynamics,and indirect effects

Despite the amount of research on the inducible defenses of prey against predators, our understanding of the long‐term significance of non‐lethal predators on prey phenotypes, prey population dynamics, and community structure has rarely been explored. Our objectives were to assess the effects of predators on prey defenses, prey population dynamics, and the relative magnitude of density‐ versus trait‐mediated indirect interactions (DMIIs and TMIIs) over multiple prey generations. Using a freshwater snail and three common snail predators, we constructed a series of community treatments with pond mesocosms that manipulated trophic structure, the identity of the top predator, and whether predators were caged or uncaged. We quantified snail phenotypes, snail population size, and resource abundance over multiple snail generations. We found that snails were expressing inducible defenses in our system although the magnitude of the responses varied over time and across predator species. Despite the expression of inducible defenses, caged predators did not reduce snail population size. There also was no evidence of TMIIs throughout the experiment suggesting that TMIIs have a minimal role in the long‐term structure of our communities. The absence of TMIIs was largely driven by the lack of predator‐induced reductions in resource consumption and the lack of consistent reductions in population size with predator cues. In contrast, we detected strong DMIIs associated with lethal predators suggesting that DMIIs are the dominant long‐term mechanism influencing community structure. Our results demonstrate that although predators can have significant effects on prey phenotypes and sometimes cause short‐term TMIIs, there may be few long‐term consequences of these responses on population dynamics and indirect interactions, at least within simple food webs. Research directed towards addressing the long‐term consequences of predator–prey interactions within communities will help to reveal whether the conclusions and predictions generated from short‐term experiments are applicable over ecological and evolutionary timescales.     

Numerical Study of Propulsion Mechanism for Oscillating Rigid and Flexible Tuna-Tails

Numerical study on the unsteady hydrodynamic characteristics of oscillating rigid and flexible tuna-tails in viscous flow-field is performed.Investigations are conducted using Reynolds-Averaged Navier-Stokes (RANS) equations with a moving adaptive mesh.The effect of swimming speed,flapping amplitude,frequency and flexure amplitude on the propulsion performance of the rigid and flexible tuna-tails are investigated.Computational results reveal that a pair of leading edge vortices develop along the tail surface as it undergoes an oscillating motion.The propulsive efficiency has a strong correlation with various locomotive parameters.Peak propulsive efficiency can be obtained by adjusting these parameters.Particularly,when input power coefficient is less than 2.8,the rigid tail generates larger thrust force and higher propulsive efficiency than flexible tail.However,when input power coefficient is larger than 2.8,flexible tail is superior to rigid tail.     

The effect of Reynolds number on the propulsive efficiency of a biomorphic pulsed-jet underwater vehicle

The effect of Reynolds number on the propulsive efficiency of pulsed-jet propulsion was studied experimentally on a self-propelled, pulsed-jet underwater vehicle, dubbed Robosquid due to the similarity of its propulsion system with squid. Robosquid was tested for jet slug length-to-diameter ratios (L/D) in the range 2-6 and dimensionless frequency (St(L)) in the range 0.2-0.6 in a glycerin-water mixture. Digital particle image velocimetry was used for measuring the impulse and energy of jet pulses from the velocity and vorticity fields of the jet flow to calculate the pulsed-jet propulsive efficiency, and compare it with an equivalent steady jet system. Robosquid's Reynolds number (Re) based on average vehicle velocity and vehicle diameter ranged between 37 and 60. The current results for propulsive efficiency were compared to the previously published results in water where Re ranged between 1300 and 2700. The results showed that the average propulsive efficiency decreased by 26% as the average Re decreased from 2000 to 50 while the ratio of pulsed-jet to steady jet efficiency (η(P)/η(P, ss)) increased up to 0.15 (26%) as the Re decreased over the same range and for similar pulsing conditions. The improved η(P)/η(P, ss) at lower Re suggests that pulsed-jet propulsion can be used as an efficient propulsion system for millimeter-scale propulsion applications. The Re = 37-60 conditions in the present investigation, showed a reduced dependence of η(P) and η(P)/η(P, ss)on L/D compared to higher Re results. This may be due to the lack of clearly observed vortex ring pinch-off as L/D increased for this Re regime.     

Nematocyst composition of the cubomedusan Chiropsalmus quadrigatuschanges with growth

The nematocysts of Chiropsalmus quadrigatus (Cubozoa; Cubomedusa; Chirodropidae) were examined to determine if their composition changes with an increase in body size. Fixed tentacles of specimens collected in Okinawa, Japan, were homogenized and their nematocysts were observed under a differential interference contrast microscope. Six nematocyst types were observed in medusae of all sizes microbasic mastigophores (MM), large and small trirhopaloids (lTR and sTR), holotrichous isorhizas (HI), ellipsoidal isorhizas (eI), and ovoid isorhizas (oI). Two other nematocysts, large ovoid isorhizas (loI) and microbasic euryteles (ME), were observed only in small individuals. There was also marked difference in proportion of tentacular nematocysts between small and large individuals. HI was the dominant type in small specimens, while MM and eI were predominant in large specimens. Nematocyst composition in the bell and pedalia also differed between small and large individuals. Bells of small medusae contained oI and sTR, while only oI were observed in most large individuals. The pedalia of small medusae had clusters of MM, ME, sTR, and oI. Such single clusters on pedalium bases were characteristic of small individuals. The pedalia of large individuals contained scattered oI. Tentacles of medusae are used for prey capture, so the changes in the major type of nematocysts in tentacles may reflect changes in prey type.