SUMMARY Flow patterns generated by medusan swimmers such as jellyfish are known to differ according the morphology of the various animal species. Oblate medusae have been previously observed to generate vortex ring structures during the propulsive cycle. Owing to the inherent physical coupling between locomotor and feeding structures in these animals, the dynamics of vortex ring formation must be robustly tuned to facilitate effective functioning of both systems. To understand how this is achieved, we employed dye visualization techniques on scyphomedusae (Aurelia aurita) observed swimming in their natural marine habitat. The flow created during each propulsive cycle consists of a toroidal starting vortex formed during the power swimming stroke, followed by a stopping vortex of opposite rotational sense generated during the recovery stroke. These two vortices merge in a laterally oriented vortex superstructure that induces flow both toward the subumbrellar feeding surfaces and downstream. The lateral vortex motif discovered here appears to be critical to the dual function of the medusa bell as a flow source for feeding and propulsion. Furthermore, vortices in the animal wake have a greater volume and closer spacing than predicted by prevailing models of medusan swimming. These effects are shown to be advantageous for feeding and swimming performance, and are an important consequence of vortex interactions that have been previously neglected.

Gelatinous zooplankton populations are well known for their ability to take over perturbed ecosystems. The ability of these animals to outcompete and functionally replace fish that exhibit an effective visual predatory mode is counterintuitive because jellyfish are described as inefficient swimmers that must rely on direct contact with prey to feed. We show that jellyfish exhibit a unique mechanism of passive energy recapture, which is exploited to allow them to travel 30% further each swimming cycle, thereby reducing metabolic energy demand by swimming muscles. By accounting for large interspecific differences in net metabolic rates, we demonstrate, contrary to prevailing views, that the jellyfish (Aurelia aurita) is one of the most energetically efficient propulsors on the planet, exhibiting a cost of transport (joules per kilogram per meter) lower than other metazoans. We estimate that reduced metabolic demand by passive energy recapture improves the cost of transport by 48%, allowing jellyfish to achieve the large sizes required for sufficient prey encounters. Pressure calculations, using both computational fluid dynamics and a newly developed method from empirical velocity field measurements, demonstrate that this extra thrust results from positive pressure created by a vortex ring underneath the bell during the refilling phase of swimming. These results demonstrate a physical basis for the ecological success of medusan swimmers despite their simple body plan. Results from this study also have implications for bioinspired design, where low-energy propulsion is required.swimming efficiency | animal-fluid interactions

We describe and characterize a method for estimating the pressure field corresponding to velocity field measurements such as those obtained by using particle image velocimetry. The pressure gradient is estimated from a time series of velocity fields for unsteady calculations or from a single velocity field for quasi-steady calculations. The corresponding pressure field is determined based on median polling of several integration paths through the pressure gradient field in order to reduce the effect of measurement errors that accumulate along individual integration paths. Integration paths are restricted to the nodes of the measured velocity field, thereby eliminating the need for measurement interpolation during this step and significantly reducing the computational cost of the algorithm relative to previous approaches. The method is validated by using numerically simulated flow past a stationary, two-dimensional bluff body and a computational model of a three-dimensional, selfpropelled anguilliform swimmer to study the effects of spatial and temporal resolution, domain size, signal-to-noise ratio and out-ofplane effects. Particle image velocimetry measurements of a freely swimming jellyfish medusa and a freely swimming lamprey are analyzed using the method to demonstrate the efficacy of the approach when applied to empirical data.

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Abstract. Medusae were the earliest animals to evolve muscle-powered swimming in the seas. Although medusae have achieved diverse and prominent ecological roles throughout the world's oceans, we argue that the primitive organization of cnidarian muscle tissue limits force production and, hence, the mechanical alternatives for swimming bell function. We use a recently developed model comparing the potential force production with the hydrodynamic requirements of jet propulsion, and conclude that jet production is possible only at relatively small bell diameters. In contrast, production of a more complex wake via what we term rowing propulsion permits much larger sizes but requires a different suite of morphological features. Analysis of morphometric data from all medusan taxa independently confirms size-dependent patterns of bell forms that correspond with model predictions. Further, morphospace analysis indicates that various lineages within the Medusozoa have proceeded along either of two evolutionary trajectories. The first alternative involved restriction of jet-propelled medusan bell diameters to small dimensions. These medusae may be either solitary individuals (characteristic of Anthomedusae and Trachymedusae) or aggregates of small individual medusan units into larger colonial forms (characteristic of the nectophores of many members of the Siphonophorae). The second trajectory involved use of rowing propulsion (characteristic of Scyphozoa and some hydromedusan lineages such as the Leptomedusae and Narcomedusae) that allows much larger bell sizes. Convergence on either of the differing propulsive alternatives within the Medusozoa has emerged via parallel evolution among different medusan lineages. The distinctions between propulsive modes have important ecological ramifications because swimming and foraging are interdependent activities for medusae. Rowing swimmers are characteristically cruising predators that select different prey types from those selected by jet-propelled medusae, which are predominantly ambush predators. These relationships indicate that the different biomechanical solutions to constraints on bell function have entailed ecological consequences that are evident in the prey selection patterns and trophic impacts of contemporary medusan lineages.Additional key words: morphology, hydrodynamics, emergent properties, plankton Medusae are a diverse array of planktonic cnidarians occupying all of the world's oceans and some freshwater habitats. The Cnidaria is an ancient clade with origins in an early radiation within the basal lineage that gave rise to the rest of the animal kingdom (Valentine 2004). Although the exact relationship between the ancient Cnidaria and the rest of the Metazoa remains unresolved, it is clear that the cellular inheritance of medusae rivals even the sponges in the restricted number of cell types available for body construction (Bonner 1965;Valentine et al. 1994). Yet, unlike sponges, medusae are characterized by the evolutionary innovation of muscle-powered motility. ...

SUMMARY Fast-swimming hydromedusan jellyfish possess a characteristic funnel-shaped velum at the exit of their oral cavity that interacts with the pulsed jets of water ejected during swimming motions. It has been previously assumed that the velum primarily serves to augment swimming thrust by constricting the ejected flow in order to produce higher jet velocities. This paper presents high-speed video and dye-flow visualizations of free-swimming Nemopsis bachei hydromedusae, which instead indicate that the time-dependent velar kinematics observed during the swimming cycle primarily serve to optimize vortices formed by the ejected water rather than to affect the speed of the ejected flow. Optimal vortex formation is favorable in fast-swimming jellyfish because,unlike the jet funnelling mechanism, it allows for the minimization of energy costs while maximizing thrust forces. However, the vortex `formation number'corresponding to optimality in N. bachei is substantially greater than the value of 4 found in previous engineering studies of pulsed jets from rigid tubes. The increased optimal vortex formation number is attributable to the transient velar kinematics exhibited by the animals. A recently developed model for instantaneous forces generated during swimming motions is implemented to demonstrate that transient velar kinematics are required in order to achieve the measured swimming trajectories. The presence of velar structures in fast-swimming jellyfish and the occurrence of similar jet-regulating mechanisms in other jet-propelled swimmers (e.g. the funnel of squid) appear to be a primary factor contributing to success of fast-swimming jetters, despite their primitive body plans.

It is generally accepted that animal-fluid interactions have shaped the evolution of animals that swim and fly. However, the functional ecological advantages associated with those adaptations are currently difficult to predict on the basis of measurements of the animal-fluid interactions. We report the identification of a robust, fluid dynamic correlate of distinct ecological functions in seven jellyfish species that represent a broad range of morphologies and foraging modes. Since the comparative study is based on properties of the vortex wake--specifically, a fluid dynamical concept called optimal vortex formation--and not on details of animal morphology or phylogeny, we propose that higher organisms can also be understood in terms of these fluid dynamic organizing principles. This enables a quantitative, physically based understanding of how alterations in the fluid dynamics of aquatic and aerial animals throughout their evolution can result in distinct ecological functions.

Abstract:Simple mechanical models emulating fish have been used recently to enable targeted study of individual factors contributing to swimming locomotion without the confounding complexity of the whole fish body. Yet, unlike these uniform models, the fish body is notable for its non-uniform material properties. In particular, flexural stiffness decreases along the fish's anterior-posterior axis. To identify the role of non-uniform bending stiffness during fish-like propulsion, we studied four foil model configurations made by adhering layers of plastic sheets to produce discrete regions of high (5.5x10 in high self-propelled speeds at low costs of transport and large thrust coefficients at relatively high efficiency. Because these metrics were not all maximized together, selection of the "best" stiffness distribution will depend on actuation constraints and performance goals. These improved models enable more detailed, accurate analyses of fish-like swimming.3