Elastic pinch biomechanisms can yield consistent launch speeds regardless of projectile mass.

Energetic trade-offs are particularly pertinent to bio-ballistic systems which impart energy to projectiles exclusively during launch. We investigated such trade-offs in the spring-propelled seeds of Loropetalum chinense, Hamamelis virginiana and Fortunearia sinensis. Using similar seed-shooting mechanisms, fruits of these confamilial plants (Hamamelidaceae) span an order of magnitude in spring and seed mass. We expected that as seed mass increases, launch speed decreases. Instead, launch speed was relatively constant regardless of seed mass. We tested if fruits shoot larger seeds by storing more elastic potential energy (PE). Spring mass and PE increased as seed mass increased (in order of increasing seed mass: L. chinense, H. virginiana, F. sinensis). As seed mass to spring mass ratio increased (ratios: H. virginiana = 0.50, F. sinensis = 0.65, L. chinense = 0.84), mass-specific PE storage increased. The conversion efficiency of PE to seed kinetic energy (KE) decreased with increasing fruit mass. Therefore, similar launch speeds across scales occurred because (i) larger fruits stored more PE and (ii) smaller fruits had higher mass-specific PE storage and improved PE to KE conversion. By examining integrated spring and projectile mechanics in our focal species, we revealed diverse, energetic scaling strategies relevant to spring-propelled systems navigating energetic trade-offs.

Mantis Shrimp Locomotion: Coordination and Variation of Hybrid Metachronal Swimming.

Across countless marine invertebrates, coordination of closely spaced swimming appendages is key to producing diverse locomotory behaviors. Using a widespread mechanism termed hybrid metachronal propulsion, mantis shrimp swim by moving five paddle-like pleopods along their abdomen in a posterior to anterior sequence during the power stroke and a near-synchronous motion during the recovery stroke. Despite the ubiquity of this mechanism, it is not clear how hybrid metachronal swimmers coordinate and modify individual appendage movements to achieve a range of swimming capabilities. Using high-speed imaging, we measured pleopod kinematics of mantis shrimp (Neogonodactylus bredini), while they performed two swimming behaviors: burst swimming and taking off from the substrate. By tracking each of the five pleopods, we tested how stroke kinematics vary across swimming speeds and the two swimming behaviors. We found that mantis shrimp achieve faster swimming speeds through a combination of higher beat frequencies, smaller stroke durations, and partially via larger stroke angles. The five pleopods exhibit non-uniform kinematics that contribute to the coordination and forward propulsion of the whole system. Micro-hook structures (retinacula) connect each of the five pleopod pairs and differ in their attachment across pleopods-possibly contributing to passive kinematic control. We compare our findings in N. bredini to previous studies to identify commonalities across hybrid metachronal swimmers at high Reynolds numbers and centimeter scales. Through our large experimental dataset and by tracking each pleopod's movements, our study reveals key parameters by which mantis shrimp adjust and control their swimming, yielding diverse locomotor abilities.

Tradeoffs explain scaling, sex differences, and seasonal oscillations in the remarkable weapons of snapping shrimp (Alpheus spp.).

Evolutionary theory suggests that individuals should express costly traits at a magnitude that optimizes the trait bearer's cost-benefit difference. Trait expression varies across a species because costs and benefits vary among individuals. For example, if large individuals pay lower costs than small individuals, then larger individuals should reach optimal cost-benefit differences at greater trait magnitudes. Using the cavitation-shooting weapons found in the big claws of male and female snapping shrimp, we test whether size- and sex-dependent expenditures explain scaling and sex differences in weapon size. We found that males and females from three snapping shrimp species (Alpheus heterochaelis, Alpheus angulosus, and Alpheus estuariensis) show patterns consistent with tradeoffs between weapon and abdomen size. For male A. heterochaelis, the species for which we had the greatest statistical power, smaller individuals showed steeper tradeoffs. Our extensive dataset in A. heterochaelis also included data about pairing, breeding season, and egg clutch size. Therefore, we could test for reproductive tradeoffs and benefits in this species. Female A. heterochaelis exhibited tradeoffs between weapon size and egg count, average egg volume, and total egg mass volume. For average egg volume, smaller females exhibited steeper tradeoffs. Furthermore, in males but not females, large weapons were positively correlated with the probability of being paired and the relative size of their pair mates. In conclusion, we identified size-dependent tradeoffs that could underlie reliable scaling of costly traits. Furthermore, weapons are especially beneficial to males and burdensome to females, which could explain why males have larger weapons than females.

From deer antlers to crab claws, weapons are some of the most elaborate and enormous structures in the animal kingdom. Within a species, weapon size generally increases with the size and condition of an individual, and those with larger weapons are usually better at fending off more diminutive competitors. Although it may seem desirable for all individuals to have large weapons, size varies greatly within a species. The 'handicap principle' proposes that the cost of bearing a weapon dictates the variation in weapon size. Smaller or less fit individuals pay more for weapons than larger or fitter animals, so smaller individuals tend to grow smaller weapons. Although popular, only a handful of studies have demonstrated experimental evidence that supports this theory. To test the handicap principle, Dinh and Patek studied a group of crustaceans known as snapping shrimp. Each shrimp has one enlarged claw that it uses as a weapon to fire imploding vapor bubbles at opponents during fights. Larger snapping shrimp have bigger enlarged claws and tend to win more contests. Males also have larger weapons than females, and this sex difference is amplified during the breeding season. Dinh and Patek studied weapon size in several species of snapping shrimp. Measurements showed that after controlling for body size, individuals with larger weapons had smaller abdomens, suggesting there is a tradeoff between weapon size and abdomen size. Furthermore, small males exhibited the steepest tradeoff, in line with the handicap principle. Snapping shrimp also showed sex-specific costs and benefits. After controlling for body size, females with larger weapons produced fewer and smaller eggs, while males with larger weapons were more likely to be paired with females and generally paired with larger females. This suggests that weapons are particularly burdensome to female shrimp and particularly beneficial to males, especially during the breeding season. These findings provide elusive evidence for the handicap principle and extend the theory to explain sex and seasonal differences in the size of snapping shrimp weapons. More broadly, the findings highlight the value of studying both male and female animal weapons when, historically, the focus has been on male weaponry.

Latch-mediated spring actuation (LaMSA): the power of integrated biomechanical systems.

Across the tree of life - from fungi to frogs - organisms wield small amounts of energy to generate fast and potent movements. These movements are propelled with elastic structures, and their loading and release are mediated by latch-like opposing forces. They comprise a class of elastic mechanisms termed latch-mediated spring actuation (LaMSA). Energy flow through LaMSA begins when an energy source loads elastic element(s) in the form of elastic potential energy. Opposing forces, often termed latches, prevent movement during loading of elastic potential energy. As the opposing forces are shifted, reduced or removed, elastic potential energy is transformed into kinetic energy of the spring and propelled mass. Removal of the opposing forces can occur instantaneously or throughout the movement, resulting in dramatically different outcomes for consistency and control of the movement. Structures used for storing elastic potential energy are often distinct from mechanisms that propel the mass: elastic potential energy is often distributed across surfaces and then transformed into localized mechanisms for propulsion. Organisms have evolved cascading springs and opposing forces not only to serially reduce the duration of energy release, but often to localize the most energy-dense events outside of the body to sustain use without self-destruction. Principles of energy flow and control in LaMSA biomechanical systems are emerging at a rapid pace. New discoveries are catalyzing remarkable growth of the historic field of elastic mechanisms through experimental biomechanics, synthesis of novel materials and structures, and high-performance robotics systems.

Developing elastic mechanisms: ultrafast motion and cavitation emerge at the millimeter scale in juvenile snapping shrimp.

Organisms such as jumping froghopper insects and punching mantis shrimp use spring-based propulsion to achieve fast motion. Studies of elastic mechanisms have primarily focused on fully developed and functional mechanisms in adult organisms. However, the ontogeny and development of these mechanisms can provide important insights into the lower size limits of spring-based propulsion, the ecological or behavioral relevance of ultrafast movement, and the scaling of ultrafast movement. Here, we examined the development of the spring-latch mechanism in the bigclaw snapping shrimp, Alpheus heterochaelis (Alpheidae). Adult snapping shrimp use an enlarged claw to produce high-speed strikes that generate cavitation bubbles. However, until now, it was unclear when the elastic mechanism emerges during development and whether juvenile snapping shrimp can generate cavitation at this size. We reared A. heterochaelis from eggs, through their larval and postlarval stages. Starting 1 month after hatching, the snapping shrimp snapping claw gradually developed a spring-actuated mechanism and began snapping. We used high-speed videography (300,000 frames s-1) to measure juvenile snaps. We discovered that juvenile snapping shrimp generate the highest recorded accelerations (5.8×105±3.3×105 m s-2) for repeated-use, underwater motion and are capable of producing cavitation at the millimeter scale. The angular velocity of snaps did not change as juveniles grew; however, juvenile snapping shrimp with larger claws produced faster linear speeds and generated larger, longer-lasting cavitation bubbles. These findings establish the development of the elastic mechanism and cavitation in snapping shrimp and provide insights into early life-history transitions in spring-actuated mechanisms.

Geometric latches enable tuning of ultrafast, spring-propelled movements.

The smallest, fastest, repeated-use movements are propelled by power-dense elastic mechanisms, yet the key to their energetic control may be found in the latch-like mechanisms that mediate transformation from elastic potential energy to kinetic energy. Here, we tested how geometric latches enable consistent or variable outputs in ultrafast, spring-propelled systems. We constructed a reduced-order mathematical model of a spring-propelled system that uses a torque reversal (over-center) geometric latch. The model was parameterized to match the scales and mechanisms of ultrafast systems, specifically snapping shrimp. We simulated geometric and energetic configurations that enabled or reduced variation of strike durations and dactyl rotations given variation of stored elastic energy and latch mediation. Then, we collected an experimental dataset of the energy storage mechanism and ultrafast snaps of live snapping shrimp (Alpheus heterochaelis) and compared our simulations with their configuration. We discovered that snapping shrimp deform the propodus exoskeleton prior to the strike, which may contribute to elastic energy storage. Regardless of the amount of variation in spring loading duration, strike durations were far less variable than spring loading durations. When we simulated this species' morphological configuration in our mathematical model, we found that the low variability of strike duration is consistent with their torque reversal geometry. Even so, our simulations indicate that torque reversal systems can achieve either variable or invariant outputs through small adjustments to geometry. Our combined experiments and mathematical simulations reveal the capacity of geometric latches to enable, reduce or enhance variation of ultrafast movements in biological and synthetic systems.

Spring and latch dynamics can act as control pathways in ultrafast systems.

Ultrafast movements propelled by springs and released by latches are thought limited to energetic adjustments prior to movement, and seemingly cannot adjust once movement begins. Even so, across the tree of life, ultrafast organisms navigate dynamic environments and generate a range of movements, suggesting unrecognized capabilities for control. We develop a framework of control pathways leveraging the non-linear dynamics of spring-propelled, latch-released systems. We analytically model spring dynamics and develop reduced-parameter models of latch dynamics to quantify how they can be tuned internally or through changing external environments. Using Lagrangian mechanics, we test feedforward and feedback control implementation via spring and latch dynamics. We establish through empirically-informed modeling that ultrafast movement can be controllably varied during latch release and spring propulsion. A deeper understanding of the interconnection between multiple control pathways, and the tunability of each control pathway, in ultrafast biomechanical systems presented here has the potential to expand the capabilities of synthetic ultra-fast systems and provides a new framework to understand the behaviors of fast organisms subject to perturbations and environmental non-idealities.

Dual spring force couples yield multifunctionality and ultrafast, precision rotation in tiny biomechanical systems.

Small organisms use propulsive springs rather than muscles to repeatedly actuate high acceleration movements, even when constrained to tiny displacements and limited by inertial forces. Through integration of a large kinematic dataset, measurements of elastic recoil, energetic math modeling and dynamic math modeling, we tested how trap-jaw ants (Odontomachus brunneus) utilize multiple elastic structures to develop ultrafast and precise mandible rotations at small scales. We found that O. brunneus develops torque on each mandible using an intriguing configuration of two springs: their elastic head capsule recoils to push and the recoiling muscle-apodeme unit tugs on each mandible. Mandibles achieved precise, planar, circular trajectories up to 49,100 rad s-1 (470,000 rpm) when powered by spring propulsion. Once spring propulsion ended, the mandibles moved with unconstrained and oscillatory rotation. We term this mechanism a 'dual spring force couple', meaning that two springs deliver energy at two locations to develop torque. Dynamic modeling revealed that dual spring force couples reduce the need for joint constraints and thereby reduce dissipative joint losses, which is essential to the repeated use of ultrafast, small systems. Dual spring force couples enable multifunctionality: trap-jaw ants use the same mechanical system to produce ultrafast, planar strikes driven by propulsive springs and for generating slow, multi-degrees of freedom mandible manipulations using muscles, rather than springs, to directly actuate the movement. Dual spring force couples are found in other systems and are likely widespread in biology. These principles can be incorporated into microrobotics to improve multifunctionality, precision and longevity of ultrafast systems.

A physical model of mantis shrimp for exploring the dynamics of ultrafast systems.

Efficient and effective generation of high-acceleration movement in biology requires a process to control energy flow and amplify mechanical power from power density-limited muscle. Until recently, this ability was exclusive to ultrafast, small organisms, and this process was largely ascribed to the high mechanical power density of small elastic recoil mechanisms. In several ultrafast organisms, linkages suddenly initiate rotation when they overcenter and reverse torque; this process mediates the release of stored elastic energy and enhances the mechanical power output of extremely fast, spring-actuated systems. Here we report the discovery of linkage dynamics and geometric latching that reveals how organisms and synthetic systems generate extremely high-acceleration, short-duration movements. Through synergistic analyses of mantis shrimp strikes, a synthetic mantis shrimp robot, and a dynamic mathematical model, we discover that linkages can exhibit distinct dynamic phases that control energy transfer from stored elastic energy to ultrafast movement. These design principles are embodied in a 1.5-g mantis shrimp scale mechanism capable of striking velocities over 26 m [Formula: see text] in air and 5 m [Formula: see text] in water. The physical, mathematical, and biological datasets establish latching mechanics with four temporal phases and identify a nondimensional performance metric to analyze potential energy transfer. These temporal phases enable control of an extreme cascade of mechanical power amplification. Linkage dynamics and temporal phase characteristics are easily adjusted through linkage design in robotic and mathematical systems and provide a framework to understand the function of linkages and latches in biological systems.

Hybrid Metachronal Rowing Augments Swimming Speed and Acceleration via Increased Stroke Amplitude.

Numerous aquatic invertebrates use drag-based metachronal rowing for swimming, in which closely spaced appendages are oscillated starting from the posterior, with each appendage phase-shifted in time relative to its neighbor. Continuously swimming species such as Antarctic krill generally use "pure metachronal rowing" consisting of a metachronal power stroke and a metachronal recovery stroke, while burst swimming species such as many copepods and mantis shrimp typically use "hybrid metachronal rowing" consisting of a metachronal power stroke followed by a synchronous or nearly synchronous recovery stroke. Burst swimming organisms need to rapidly accelerate in order to capture prey and/or escape predation, and it is unknown whether hybrid metachronal rowing can augment acceleration and swimming speed compared to pure metachronal rowing. Simulations of rigid paddles undergoing simple harmonic motion showed that collisions between adjacent paddles restrict the maximum stroke amplitude for pure metachronal rowing. Hybrid metachronal rowing similar to that observed in mantis shrimp (Neogonodactylus bredini) permits oscillation at larger stroke amplitude while avoiding these collisions. We comparatively examined swimming speed, acceleration, and wake structure of pure and hybrid metachronal rowing strategies by using a self-propelling robot. Both swimming speed and peak acceleration of the robot increased with increasing stroke amplitude. Hybrid metachronal rowing permitted operation at larger stroke amplitude without collision of adjacent paddles on the robot, augmenting swimming speed and peak acceleration. Hybrid metachronal rowing generated a dispersed wake unlike narrower, downward-angled jets generated by pure metachronal rowing. Our findings suggest that burst swimming animals with small appendage spacing, such as copepods and mantis shrimp, can use hybrid metachronal rowing to generate large accelerations via increasing stroke amplitude without concern of appendage collision.

Scaling and development of elastic mechanisms: the tiny strikes of larval mantis shrimp.

Latch-mediated spring actuation (LaMSA) is used by small organisms to produce high acceleration movements. Mathematical models predict that acceleration increases as LaMSA systems decrease in size. Adult mantis shrimp use a LaMSA mechanism in their raptorial appendages to produce extremely fast strikes. Until now, however, it was unclear whether mantis shrimp at earlier life-history stages also strike using elastic recoil and latch mediation. We tested whether larval mantis shrimp (Gonodactylaceus falcatus) use LaMSA and, because of their smaller size, achieve higher strike accelerations than adults of other mantis shrimp species. Based on microscopy and kinematic analyses, we discovered that larval G. falcatus possess the components of, and actively use, LaMSA during their fourth larval stage, which is the stage of development when larvae begin feeding. Larvae performed strikes at high acceleration and speed (mean: 4.133×105 rad s-2, 292.7 rad s-1; 12 individuals, 25 strikes), which are of the same order of magnitude as for adults - even though adult appendages are up to two orders of magnitude longer. Larval strike speed (mean: 0.385 m s-1) exceeded the maximum swimming speed of similarly sized organisms from other species by several orders of magnitude. These findings establish the developmental timing and scaling of the mantis shrimp LaMSA mechanism and provide insights into the kinematic consequences of scaling limits in tiny elastic mechanisms.

Snaps of a tiny amphipod push the boundary of ultrafast, repeatable movement.

Surprisingly, the fastest motions are not produced by large animals or robots. Rather, small organisms or structures, including cnidarian stinging cells, fungal shooting spores, and mandible strikes of ants, termites, and spiders, hold the world acceleration records.1-5 These diverse systems share common features: they rapidly convert potential energy - stored in deformed material or fluid - into kinetic energy when a latch is released.4-6 However, the fastest of these are not repeatable, because mechanical components are broken or ejected.5,6 Furthermore, some of these systems must overcome the added challenge of moving in water, where high density and viscosity constrain acceleration at small sizes. Here we report the kinematics of repeatable, ultrafast snaps by tiny marine amphipods (Dulichiella cf. appendiculata). Males use their enlarged major claw, which can exceed 30% of body mass, to snap a 1 mm-long dactyl with a diameter equivalent to a human hair (184 µm). The claw snaps closed extremely rapidly, averaging 93 µs, 17 m s-1, and 2.4 x 105 m s-2. These snaps are among the smallest and fastest of any documented repeatable movement, and are sufficiently fast to operate in the inertial hydrodynamic regime (Reynolds number (Re) >10,000). They generate audible pops and rapid water jets, which occasionally yield cavitation, and may be used for defense. These amphipod snaps push the boundaries of acceleration and size for repeatable movements, particularly in water, and exemplify how new biomechanical insights can arise from unassuming animals. VIDEO ABSTRACT.

Pendulum-based measurements reveal impact dynamics at the scale of a trap-jaw ant.

Small organisms can produce powerful, sub-millisecond impacts by moving tiny structures at high accelerations. We developed and validated a pendulum device to measure the impact energetics of microgram-sized trap-jaw ant mandibles accelerated against targets at 105 m s-2 Trap-jaw ants (Odontomachus brunneus; 19 individuals, 212 strikes) were suspended on one pendulum and struck swappable targets that were either attached to an opposing pendulum or fixed in place. Mean post-impact kinetic energy (energy from a strike converted to pendulum motion) was higher with a stiff target (21.0-21.5 µJ) than with a compliant target (6.4-6.5 µJ). Target mobility had relatively little influence on energy transfer. Mean contact duration of strikes against stiff targets was shorter (3.9-4.5 ms) than against compliant targets (6.2-7.9 ms). Shorter contact duration was correlated with higher post-impact kinetic energy. These findings contextualize and provide an energetic explanation for the diverse, natural uses of trap-jaw ant strikes such as impaling prey, launching away threats and performing mandible-powered jumps. The strong effect of target material on energetic exchange suggests material interactions as an avenue for tuning performance of small, high acceleration impacts. Our device offers a foundation for novel research into the ecomechanics and evolution of tiny biological impacts and their application in synthetic systems.

Latch-based control of energy output in spring actuated systems.

The inherent force-velocity trade-off of muscles and motors can be overcome by instead loading and releasing energy in springs to power extreme movements. A key component of this paradigm is the latch that mediates the release of spring energy to power the motion. Latches have traditionally been considered as switches; they maintain spring compression in one state and allow the spring to release energy without constraint in the other. Using a mathematical model of a simplified contact latch, we reproduce this instantaneous release behaviour and also demonstrate that changing latch parameters (latch release velocity and radius) can reduce and delay the energy released by the spring. We identify a critical threshold between instantaneous and delayed release that depends on the latch, spring, and mass of the system. Systems with stiff springs and small mass can attain a wide range of output performance, including instantaneous behaviour, by changing latch release velocity. We validate this model in both a physical experiment as well as with data from the Dracula ant, Mystrium camillae, and propose that latch release velocity can be used in both engineering and biological systems to control energy output.

The effect of size-scale on the kinematics of elastic energy release.

Elastically-driven motion has been used as a strategy to achieve high speeds in small organisms and engineered micro-robotic devices. We examine the size-scaling relations determining the limit of elastic energy release from elastomer bands that efficiently cycle mechanical energy with minimal loss. The maximum center-of-mass velocity of the elastomer bands was found to be size-scale independent, while smaller bands demonstrated larger accelerations and shorter durations of elastic energy release. Scaling relationships determined from these measurements are consistent with the performance of small organisms and engineered devices which utilize elastic elements to power motion.

Adhesive latching and legless leaping in small, worm-like insect larvae.

Jumping is often achieved using propulsive legs, yet legless leaping has evolved multiple times. We examined the kinematics, energetics and morphology of long-distance jumps produced by the legless larvae of gall midges (Asphondylia sp.). They store elastic energy by forming their body into a loop and pressurizing part of their body to form a transient 'leg'. They prevent movement during elastic loading by placing two regions covered with microstructures against each other, which likely serve as a newly described adhesive latch. Once the latch releases, the transient 'leg' launches the body into the air. Their average takeoff speeds (mean: 0.85 m s-1; range: 0.39-1.27 m s-1) and horizontal travel distances (up to 36 times body length or 121 mm) rival those of legged insect jumpers and their mass-specific power density (mean: 910 W kg-1; range: 150-2420 W kg-1) indicates the use of elastic energy storage to launch the jump. Based on the forces reported for other microscale adhesive structures, the adhesive latching surfaces are sufficient to oppose the loading forces prior to jumping. Energetic comparisons of insect larval crawling versus jumping indicate that these jumps are orders of magnitude more efficient than would be possible if the animals had crawled an equivalent distance. These discoveries integrate three vibrant areas in engineering and biology - soft robotics, small, high-acceleration systems, and adhesive systems - and point toward a rich, and as-yet untapped area of biological diversity of worm-like, small, legless jumpers.

Beyond power amplification: latch-mediated spring actuation is an emerging framework for the study of diverse elastic systems.

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.

The Power of Mantis Shrimp Strikes: Interdisciplinary Impacts of an Extreme Cascade of Energy Release.

In the course of a single raptorial strike by a mantis shrimp (Stomatopoda), the stages of energy release span six to seven orders of magnitude of duration. To achieve their mechanical feats of striking at the outer limits of speeds, accelerations, and impacts among organisms, they use a mechanism that exemplifies a cascade of energy release-beginning with a slow and forceful, spring-loading muscle contraction that lasts for hundreds of milliseconds and ending with implosions of cavitation bubbles that occur in nanoseconds. Mantis shrimp use an elastic mechanism built of exoskeleton and controlled with a latching mechanism. Inspired by both their mechanical capabilities and evolutionary diversity, research on mantis shrimp strikes has provided interdisciplinary and fundamental insights to the fields of elastic mechanisms, fluid dynamics, evolutionary dynamics, contest dynamics, the physics of fast, small systems, and the rapidly-expanding field of bioinspired materials science. Even with these myriad connections, numerous discoveries await, especially in the arena of energy flow through materials actuating and controlling fast, impact fracture resistant systems.