Although members of the Rhopalonematidae family (Cnidaria, Hydrozoa, Trachymedusae) are known to exhibit unusually powerful jet swimming in addition to their more normal slow swimming behaviour, for the most part reports are rare and anecdotal. Many species are found globally at depths of 600 - 2000 m and so observation and collection depends on using remotely operated submersible vehicles. With a combination of in-situ video footage and laboratory measurements, we have quantified kinematic aspects of this dual swimming motion and its electrophysiology. The species included are from two Rhopalonematidae clades; they are Colobonema sericeum, Pantachogon haeckeli, Crossota millsae and two species of Benthocodon. Comparison is made with Aglantha digitale, a species from a third Rhopalonematidae clade brought to the surface by natural water movement. We find that although all Rhopalonematidae appear to have two swimming modes there are marked differences in their neural anatomy, kinematics and physiology. Giant motor axons, known to conduct impulses during fast swimming in A. digitale, are absent from C. sericeum and P. haeckeli . Slow swimming is also different; in C. sericeum and its relatives it is driven by contractions restricted to the base of the bell. These behavioural differences are related to the position of the different clades on a ribosomal DNA-based phylogenetic tree. This finding allows us to pinpoint the phylogenetic branch point leading to the appearance of giant motor axons and escape swimming. They place the remarkable dual swimming behaviour of members of the Rhopalonematidae family into an evolutionary context.

ABSTRACT Intraoral food processing mechanisms are known for all major vertebrate groups, but the form and function of systems used to crush, grind or puncture food items can differ substantially between and within groups. Most vertebrates display flexible mechanisms of intraoral food processing with respect to different environmental conditions or food types. It has recently been shown that newts use cyclical loop-motions of the tongue to rasp prey against the palatal dentition. However, it remains unknown whether newts can adjust their food processing behavior in response to different food types or environmental conditions. Newts are interesting models for studying the functional adaptation to different conditions because of their unique and flexible lifestyle: they seasonally change between aquatic and terrestrial habitats, adapt their prey-capture mode to the respective environment, and consume diverse food types with different mechanical properties. Using X-ray high-speed recordings, anatomical investigations, behavioral analyses and mechanical property measurements, we tested the effects of the medium in which feeding occurs (water/air) and the food type (maggot, earthworm, cricket) on the processing behavior in Triturus carnifex . We discovered that food processing, by contrast to prey capture, differed only slightly between aquatic and terrestrial habitats. However, newts adjusted the number of processing cycles to different prey types: while maggots were processed extensively, earthworm pieces were barely processed at all. We conclude that, in addition to food mechanical properties, sensory feedback such as smell and taste appear to induce flexible processing responses, while the medium in which feeding occurs appears to have less of an effect.

ABSTRACT Many walking insects use vision for long-distance navigation, but the influence of vision on rapid walking performance that requires close-range obstacle detection and directing the limbs towards stable footholds remains largely untested. We compared Argentine ant ( Linepithema humile ) workers in light versus darkness while traversing flat and uneven terrain. In darkness, ants reduced flat-ground walking speeds by only 5%. Similarly, the approach speed and time to cross a step obstacle were not significantly affected by lack of lighting. To determine whether tactile sensing might compensate for vision loss, we tracked antennal motion and observed shifts in spatiotemporal activity as a result of terrain structure but not illumination. Together, these findings suggest that vision does not impact walking performance in Argentine ant workers. Our results help contextualize eye variation across ants, including subterranean, nocturnal and eyeless species that walk in complete darkness. More broadly, our findings highlight the importance of integrating vision, proprioception and tactile sensing for robust locomotion in unstructured environments.

ABSTRACT Movements of ectotherms are constrained by their body temperature owing to the effects of temperature on muscle physiology. As physical performance often affects the outcome of predator–prey interactions, environmental temperature can influence the ability of ectotherms to capture prey and/or defend themselves against predators. However, previous research on the kinematics of ectotherms suggests that some species may use elastic storage mechanisms when attacking or defending, thereby mitigating the effects of sub-optimal temperature. Rattlesnakes ( Crotalus spp.) are a speciose group of ectothermic viperid snakes that rely on crypsis, rattling and striking to deter predators. We examined the influence of body temperature on the behavior and kinematics of two rattlesnake species ( Crotalus oreganus helleri and Crotalus scutulatus ) when defensively striking towards a threatening stimulus. We recorded defensive strikes at body temperatures ranging from 15–35°C. We found that strike speed and speed of mouth gaping during the strike were positively correlated with temperature. We also found a marginal effect of temperature on the probability of striking, latency to strike and strike outcome. Overall, warmer snakes are more likely to strike, strike faster, open their mouth faster and reach maximum gape earlier than colder snakes. However, the effects of temperature were less than would be expected for purely muscle-driven movements. Our results suggest that, although rattlesnakes are at a greater risk of predation at colder body temperatures, their decrease in strike performance may be mitigated to some extent by employing mechanisms in addition to skeletal muscle contraction (e.g. elastic energy storage) to power strikes.

ABSTRACT Most non-mammal tetrapods have a hinge-like jaw operation restricted to vertical opening and closing movements. Many mammal jaw joints, by contrast, operate in more complex, three-dimensional (3D) ways, involving not only vertical but also propalinal (rostro-caudal) and transverse (lateral) movements. Data on intraoral food processing in lissamphibians and sauropsids has prompted a generally accepted view that these groups mostly swallow food unreduced, and that in those cases where lissamphibians and sauropsids chew, they mostly use simple vertical jaw movements for food processing. The exception to this generally accepted view is the occurrence of some propalinal chewing in sauropsids. We combined 3D kinematics and morphological analyses from biplanar high-speed video fluoroscopy and micro-computed tomography to determine how the paedomorphic salamander Siren intermedia treats captured food. We discovered not only that S. intermedia uses intraoral food processing but also that the elaborated morphology of its jaw joint facilitates mandibular motions in all three planes, resulting in complex 3D chewing. Thus, our data challenge the commonly held view that complex 3D chewing movements are exclusive to mammals, by suggesting that such mechanisms might have evolved early in the tetrapod evolution.

ABSTRACT Mantis shrimp strikes are one of the fastest animal movements, despite their occurrence in a water medium with viscous drag. Since the strike is produced by a latch-mediated spring-actuated system and not directly driven by muscle action, we predicted that strikes performed in air would be faster than underwater as a result of reduction in the medium's drag. Using high-speed video analysis of stereotyped strikes elicited from Squilla mantis , we found the exact opposite: strikes are much slower and less powerful in air than in water. S. mantis strikes in air have a similar mass and performance to latch-mediated spring-actuated jumps in locusts, suggesting a potential threshold for the energetics of a 1–2 g limb rotating in air. Drag forces induced by the media may be a key feature in the evolution of mantis shrimp strikes and provide a potential target for probing the braking system of these extremely fast movements.

ABSTRACT Controlled landing requires preparation. Mammals and bipedal birds vary how they prepare for landing by predicting the timing and magnitude of impact from the integration of visual and non-visual information. Here, we explore how the cane toad Rhinella marina – an animal that moves primarily through hopping – integrates sensory information to modulate landing preparation. Earlier work suggests that toads may modulate landing preparation using predictions of impact timing and/or magnitude based on non-visual sensory feedback during takeoff rather than visual cues about the landing itself. We disentangled takeoff and landing conditions by hopping toads off platforms of different heights while measuring electromyographic (EMG) activity of an elbow extensor (m. anconeus) and capturing high-speed images to quantify whole body and forelimb kinematics. This enabled us to test how toads integrate visual and non-visual information in landing preparation. We asked two questions: (1) when they conflict, do toads correlate landing preparation with takeoff or landing conditions? And (2) for hops with the same takeoff conditions, does visual information alter the timing of landing preparation? We found that takeoff conditions are a better predictor of the onset of landing preparation than landing conditions, but that visual information is not ignored. When hopping off higher platforms, toads start to prepare for landing later when takeoff conditions are invariant. This suggests that, unlike mammals, toads prioritize non-visual sensory feedback about takeoff conditions to coordinate landing, but that they do integrate visual information to fine-tune landing preparation.

ABSTRACT Ants are famous in the animal kingdom for their amazing load-carrying performance. Yet, the mechanisms that allow these insects to maintain their stability when carrying heavy loads have been poorly investigated. Here, we present a study of the kinematics of unloaded and loaded locomotion in the polymorphic seed-harvesting ant Messor barbarus . In this species, large ants have larger heads relative to their size than small ants. Hence, their center of mass is shifted forward, and even more so when they are carrying a load in their mandibles. We tested the hypothesis that this could lead to large ants being less statically stable than small ants, thus explaining their lower load-carrying ability. We found that large ants were indeed less statically stable than small ants when walking unloaded, but they were nonetheless able to adjust their stepping pattern to partly compensate for this instability. When ants were walking loaded on the other hand, there was no evidence of different locomotor behaviors in individuals of different sizes. Loaded ants, whatever their size, move too slowly to maintain their balance through dynamic stability. Rather, they seem to do so by clinging to the ground with their hind legs during part of a stride. We show through a straightforward model that allometric relationships have a minor role in explaining the differences in load-carrying ability between large ants and small ants, and that a simple scale effect is sufficient to explain these differences.

ABSTRACT Closed kinematic chains (CKCs), links connected to form one or more closed loops, are used as simple models of musculoskeletal systems (e.g. the four-bar linkage). Previous applications of CKCs have primarily focused on biomechanical systems with rigid links and permanently closed chains, which results in constant mobility (the total degrees of freedom of a system). However, systems with non-rigid elements (e.g. ligaments and muscles) and that alternate between open and closed chains (e.g. standing on one foot versus two) can also be treated as CKCs with changing mobility. Given that, in general, systems that have fewer degrees of freedom are easier to control, what implications might such dynamic changes in mobility have for motor control? Here, I propose a CKC classification to explain the different ways in which mobility of musculoskeletal systems can change dynamically during behavior. This classification is based on the mobility formula, taking into account the number of loops in the CKC and the nature of the constituent joint mobilities. I apply this mobility-based classification to five biomechanical systems: the human lower limbs, the operculum–lower jaw mechanism of fishes, the upper beak rotation mechanism of birds, antagonistic muscles at the human ankle joint and the human jaw processing a food item. I discuss the implications of this classification, including that mobility itself may be dynamically manipulated to simplify motor control. The principal aim of this Commentary is to provide a framework for quantifying mobility across diverse musculoskeletal systems to evaluate its potentially key role in motor control.

ABSTRACT The study of fish escape responses has provided important insights into the accelerative motions and fast response times of these animals. In addition, the accessibility of the underlying neural circuits has made the escape response a fundamental model in neurobiology. Fish escape responses were originally viewed as highly stereotypic all-or-none behaviours. However, research on a wide variety of species has shown considerable taxon-specific and context-dependent variability in the kinematics and neural control of escape. In addition, escape-like motions have been reported: these resemble escape responses kinematically, but occur in situations that do not involve a response to a threatening stimulus. This Review focuses on the diversity of escape responses in fish by discussing recent work on: (1) the types of escape responses as defined by kinematic analysis (these include C - and S -starts, and single- versus double-bend responses); (2) the diversity of neuromuscular control; (3) the variability of escape responses in terms of behaviour and kinematics within the context of predator−prey interactions; and (4) the main escape-like motions observed in various species. Here, we aim to integrate recent knowledge on escape responses and highlight rich areas for research. Rapidly developing approaches for studying the kinematics of swimming motion both in the lab and within the natural environment provide new avenues for research on these critical and common behaviours.

ABSTRACT Rapid biological movements, such as the extraordinary strikes of mantis shrimp and accelerations of jumping insects, have captivated generations of scientists and engineers. These organisms store energy in elastic structures (e.g. springs) and then rapidly release it using latches, such that movement is driven by the rapid conversion of stored elastic to kinetic energy using springs, with the dynamics of this conversion mediated by latches. Initially drawn to these systems by an interest in the muscle power limits of small jumping insects, biologists established the idea of power amplification, which refers both to a measurement technique and to a conceptual framework defined by the mechanical power output of a system exceeding muscle limits. However, the field of fast elastically driven movements has expanded to encompass diverse biological and synthetic systems that do not have muscles – such as the surface tension catapults of fungal spores and launches of plant seeds. Furthermore, while latches have been recognized as an essential part of many elastic systems, their role in mediating the storage and release of elastic energy from the spring is only now being elucidated. Here, we critically examine the metrics and concepts of power amplification and encourage a framework centered on latch-mediated spring actuation (LaMSA). We emphasize approaches and metrics of LaMSA systems that will forge a pathway toward a principled, interdisciplinary field.

ABSTRACT Most fish species use fast starts to escape from predators. Zebrafish larvae perform effective fast starts immediately after hatching. They use a C -start, where the body curls into a C -shape, and then unfolds to accelerate. These escape responses need to fulfil a number of functional demands, under the constraints of the fluid environment and the larva's body shape. Primarily, the larvae need to generate sufficient escape speed in a wide range of possible directions, in a short-enough time. In this study, we examined how the larvae meet these demands. We filmed fast starts of zebrafish larvae with a unique five-camera setup with high spatiotemporal resolution. From these videos, we reconstructed the 3D swimming motion with an automated method and from these data calculated resultant hydrodynamic forces and, for the first time, 3D torques. We show that zebrafish larvae reorient mostly in the first stage of the start by producing a strong yaw torque, often without using the pectoral fins. This reorientation is expressed as the body angle, a measure that represents the rotation of the complete body, rather than the commonly used head angle. The fish accelerates its centre of mass mostly in stage 2 by generating a considerable force peak while the fish ‘unfolds’. The escape direction of the fish correlates strongly with the amount of body curvature in stage 1, while the escape speed correlates strongly with the duration of the start. This may allow the fish to independently control the direction and speed of the escape.

ABSTRACT Many species of fish process their prey with cyclic jaw motions that grossly resemble those seen in mammalian mastication, despite starkly different tooth and jaw morphologies. The degree of similarity between the processing behaviors of these disparate taxa has implications for our understanding of convergence in vertebrate feeding systems. Here, we used XROMM (X-ray reconstruction of moving morphology) to investigate prey processing behavior of Potamotrygon motoro , the ocellate river stingray, which has recently been found to employ asymmetrical, shearing jaw motions to break down its prey. We found that P. motoro modulates its feeding kinematics to produce two distinct types of chew cycles: compressive cycles and overbite cycles. The latter are characterized by over-rotation of the upper jaw relative to the lower jaw, past the expected occlusal limit, and higher levels of bilateral asymmetry as compared with compressive chews. We did not find evidence of the mediolateral shearing motions typical of mammalian mastication, but overbite cycles appear to shear the prey item between the upper and lower toothplates in a propalinal fashion. Additionally, comparison of hyomandibular and jaw motions demonstrates that the angular cartilages decouple jaw displacement from hyomandibular displacement in rostrocaudal and mediolateral directions. The multiple similarities between mammalian mastication and the dynamic processing behavior of P. motoro support the use of sub-family Potamotrygoninae as a model for studying evolutionary convergence of mastication-like processing.

ABSTRACT Pursuit is a common behavior exhibited by animals chasing prey, competitors and potential mates. Because of their speed and maneuverability, dragonflies are frequently studied as a model system for biological pursuit. Most quantitative studies have focused on prey pursuits in captive environments. To determine whether a different pursuit strategy is used when chasing conspecifics of nearly equal speed and agility, we recorded 3D flight trajectories from nine territorial chases between male Erythemis simplicicollis dragonflies in natural field conditions. During chases, dragonflies used an interception strategy with an unusually high-magnitude gain ( k =−10.03 s −1 horizontal; −8.86 s −1 vertical) and short time delay (τ=50 ms). The product k τ determines how aggressively a pursuer corrects course to achieve interception. Previous studies of prey pursuit have found k τ values close to −1/ e (−0.37), the time-optimal value for achieving pursuit without overshooting. However, we found that dragonflies chasing conspecifics use more negative k τ (−0.50 horizontal; −0.44 vertical), resulting in pursuits with a high degree of overshooting (i.e. moving past the target and alternating position from side to side). We confirmed via simulation that the observed gain and delay produce overshooting. We propose that overshooting is an adaptive feature of conspecific chases that can be achieved with only slight modification of the strategy used for intercepting prey. Overshooting might help avoid potentially damaging collisions while exhibiting the pursuing animal's flight performance and competitive ability. Repeated close approaches might also evoke evasive responses from the other dragonfly, effectively herding the competitor out of the territory.

ABSTRACT Biologists commonly visualize different features of an organism using distinct sources of illumination. Such multichannel imaging has largely not been applied to behavioral studies because of the challenges posed by a moving subject. We address this challenge with the technique of multichannel stroboscopic videography (MSV), which synchronizes multiple strobe lights with video exposures of a single camera. We illustrate the utility of this approach with kinematic measurements of a walking cockroach ( Gromphadorhina portentosa ) and calculations of the pressure field around a swimming fish ( Danio rerio ). In both, transmitted illumination generated high-contrast images of the animal's body in one channel. Other sources of illumination were used to visualize the points of contact for the feet of the cockroach and the water flow around the fish in separate channels. MSV provides an enhanced potential for high-throughput experimentation and the capacity to integrate changes in physiological or environmental conditions in freely-behaving animals.

ABSTRACT Predatory fishes avoid detection by prey through a stealthy approach, followed by a rapid and precise fast-start strike. Although many first-feeding fish larvae strike at non-evasive prey using an S -start, the clownfish Amphiprion ocellaris feeds on highly evasive calanoid copepods from a J -shaped position, beginning 1 day post-hatch (dph). We quantified this unique strike posture by observing successful predatory interactions between larval clownfish (1 to 14 dph) and three developmental stages of the calanoid copepod Bestiolina similis . The J -shaped posture of clownfish became less tightly curled (more L -shaped) during larval development. Larvae were also less tightly curled when targeting adult copepods, which are more evasive than younger copepod stages. Strike performance measured as time to capture and as peak speed improved only slightly with larval age. Therefore, the J -posture may allow first-feeding larvae to minimize disturbance during their approach of sensitive prey, and may represent an alternative predatory strategy to the prototypical S -start.

ABSTRACT During walking, the leg motor system must continually adjust to changes in mechanical conditions, such as the inclination of the ground. To understand the underlying control, it is important to know how changes in leg muscle activity relate to leg kinematics (movements) and leg dynamics (forces, torques). Here, we studied these parameters in hindlegs of stick insects ( Carausius morosus ) during level and uphill/downhill (±45 deg) walking, using a combination of electromyography, 3D motion capture and ground reaction force measurements. We find that some kinematic parameters including leg joint angles and body height vary across walking conditions. However, kinematics vary little compared with dynamics: horizontal leg forces and torques at the thorax–coxa joint (leg protraction/retraction) and femur–tibia joint (leg flexion/extension) tend to be stronger during uphill walking and are reversed in sign during downhill walking. At the thorax–coxa joint, the different mechanical demands are met by adjustments in the timing and magnitude of antagonistic muscle activity. Adjustments occur primarily in the first half of stance after the touch-down of the leg. When insects transition from level to incline walking, the characteristic adjustments in muscle activity occur with the first step of the leg on the incline, but not in anticipation. Together, these findings indicate that stick insects adjust leg muscle activity on a step-by-step basis so as to maintain a similar kinematic pattern under different mechanical demands. The underlying control might rely primarily on feedback from leg proprioceptors signaling leg position and movement.

ABSTRACT Characterisation of an organism's performance in different habitats provides insight into the conditions that allow it to survive and reproduce. In recent years, the northern quoll ( Dasyurus hallucatus ) – a medium-sized semi-arboreal marsupial native to northern Australia – has undergone significant population declines within open forest, woodland and riparian habitats, but less so in rocky areas. To help understand this decline, we quantified the biomechanical performance of wild northern quolls as they ran up inclined narrow (13 mm pole) and inclined wide (90 mm platform) substrates. We predicted that quolls may possess biomechanical adaptations to increase stability on narrow surfaces, which are more common in rocky habitats. Our results showed that quolls have some biomechanical characteristics consistent with a stability advantage on narrow surfaces. This includes the coupled use of limb pairs, as indicated via a decrease in footfall time, and an ability to produce corrective torques to counteract the toppling moments commonly encountered during gait on narrow surfaces. However, speed was constrained on narrow surfaces, and quolls did not adopt diagonal sequence gaits, unlike true arboreal specialists such as primates. In comparison with key predators, such as cats and dogs, northern quolls appear inferior in terrestrial environments but have a stability advantage at higher speeds on narrow supports. This may partially explain the heterogeneous declines in northern quoll populations among various habitats on mainland Australia.

ABSTRACT Animals use diverse feeding strategies to capture and consume prey, with many species switching between strategies to accommodate different prey. Many marine animals exhibit behavioral flexibility when feeding to deal with spatial and temporal heterogeneity in prey resources. However, little is known about flexibility in the feeding behavior of many large marine predators. Here, we documented the feeding behavior and kinematics of the endangered Hawaiian monk seal ( Neomonachus schauinslandi , n =7) through controlled feeding trials. Seals were fed multiple prey types (e.g. night smelt, capelin, squid and herring) that varied in size and shape to examine behavioral flexibility in feeding. Hawaiian monk seals primarily used suction feeding (91% of all feeding trials) across all prey types, but biting, specifically pierce feeding, was also observed (9% of all feeding trials). Suction feeding was characterized by shorter temporal events, a smaller maximum gape and gape angle, and a fewer number of jaw motions than pierce feeding; suction feeding kinematic performance was also more variable compared with pierce feeding. Seals showed behavioral flexibility in their use of the two strategies. Suction feeding was used most frequently when targeting small to medium sized prey and biting was used with increasing frequency on larger prey. The feeding kinematics differed between feeding strategies and prey types, showing that Hawaiian monk seals adjusted their behaviors to particular feeding contexts. Hawaiian monk seals are opportunistic marine predators and their ability to adapt their feeding strategy and behavior to specific foraging scenarios allows them to target diverse prey resources.

ABSTRACT Crocodilians have played a significant role in evolutionary studies of archosaurs. Given that several major shifts in forelimb function occur within Archosauria, forelimb morphologies of living crocodilians are of particular importance in assessing locomotor evolutionary scenarios. A previous X-ray investigation of walking alligators revealed substantial movement of the shoulder girdle, but as the sternal cartilages do not show up in X-ray, the source of the mobility could not be conclusively determined. Scapulocoracoid movement was interpreted to indicate independent sliding of each coracoid at the sternocoracoid joint; however, rotations of the sternum could also produce similar displacement of the scapulocoracoids. Here, we present new data employing marker-based XROMM (X-ray reconstruction of moving morphology), wherein simultaneous biplanar X-ray video and surgically implanted radio-opaque markers permit precise measurement of the vertebral axis, sternum and coracoid in walking alligators. We found that movements of the sternum and sternocoracoid joint both contribute to shoulder girdle mobility and stride length, and that the sternocoracoid contribution was less than previously estimated. On average, the joint contributions to stride length (measured with reference to a point on the distal radius, thus excluding wrist motion) are as follows: thoracic vertebral rotation 6.2±3.7%, sternal rotation 11.1±2.5%, sternocoracoid joint 10.1±5.2%, glenohumeral joint 40.1±7.8% and elbow 31.1±4.2%. To our knowledge, this is the first evidence of sternal movement relative to the vertebral column (presumably via rib joints) contributing to stride length in tetrapods.