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Free, publicly-accessible full text available May 19, 2026
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Free, publicly-accessible full text available May 19, 2026
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ABSTRACT Centipedes coordinate body and limb flexion to generate propulsion. On flat, solid surfaces, the limb-stepping patterns can be characterized according to the direction in which limb-aggregates propagate, opposite to (retrograde) or with the direction of motion (direct). It is unknown how limb and body dynamics are modified in terrain with terradynamic complexity more representative of these animal's natural heterogeneous environments. Here, we investigated how centipedes that use retrograde and direct limb-stepping patterns, Scolopendra polymorpha and Scolopocryptops sexspinosus, respectively, coordinate their body and limbs to navigate laboratory environments which present footstep challenges and terrain rugosity. We recorded the kinematics and measured the locomotive performance of these animals traversing two rugose terrains with randomly distributed step heights and compared the kinematics with those on a flat frictional surface. Scolopendra polymorpha exhibited similar body and limb dynamics across all terrains and a decrease in speed with increased terrain rugosity. Unexpectedly, when placed in a rugose terrain, S. sexspinosus changed the direction of the limb-stepping pattern from direct to retrograde. Further, for both species, traversal of these rugose terrains was facilitated by hypothesized passive mechanics: upon horizontal collision of a limb with a block, the limb bent and later continued the stepping pattern. Although centipedes have many degrees of freedom, our results suggest these animals negotiate limb–substrate interactions and navigate complex terrains leveraging the innate flexibility of their limbs to simplify control.more » « less
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Locomotion is typically studied either in continuous media where bodies and legs experience forces generated by the flowing medium or on solid substrates dominated by friction. In the former, centralized whole-body coordination is believed to facilitate appropriate slipping through the medium for propulsion. In the latter, slip is often assumed minimal and thus avoided via decentralized control schemes. We find in laboratory experiments that terrestrial locomotion of a meter-scale multisegmented/legged robophysical model resembles undulatory fluid swimming. Experiments varying waves of leg stepping and body bending reveal how these parameters result in effective terrestrial locomotion despite seemingly ineffective isotropic frictional contacts. Dissipation dominates over inertial effects in this macroscopic-scaled regime, resulting in essentially geometric locomotion on land akin to microscopic-scale swimming in fluids. Theoretical analysis demonstrates that the high-dimensional multisegmented/legged dynamics can be simplified to a centralized low-dimensional model, which reveals an effective resistive force theory with an acquired viscous drag anisotropy. We extend our low-dimensional, geometric analysis to illustrate how body undulation can aid performance in non–flat obstacle-rich terrains and also use the scheme to quantitatively model how body undulation affects performance of biological centipede locomotion (the desert centipede Scolopendra polymorpha ) moving at relatively high speeds (∼0.5 body lengths/sec). Our results could facilitate control of multilegged robots in complex terradynamic scenarios.more » « less
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Terrain irregularities in natural environments present mobility challenges for autonomous robots and vehicles. Loosely consolidated sandy slopes flow unpredictably when perturbed, often leading to locomotion failure. Systematic experiments with various robot morphologies on flowable terrains feature open‐loop quasistatic gait strategies that remodel the terrain to aid locomotor kinematics. On a sloped terrain of granular media near the critical angle, a laboratory‐scale rover robot induces a flow via a localized fluidization gait to remodel local terrain and succeed in locomotion. A Bayesian optimization machine learning approach that modulates this gait strategy then finds a pattern of selectively fluidizing and solidifying terrain to climb slopes rapidly. In a biped walker robot, a cleated foot design dynamically manipulates the stress fields of flowable slopes. The deeply submerged cleats remodel the shear response of the material by creating jammed regions behind them which then improve forward progression by reducing slip when compared to a flat foot. The “robophysics” approach of systematic experiments exploring terrain reconfiguration combined with future machine learning models of flowable terrain evolution can augment gait discovery for future robots.more » « less
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null (Ed.)Abstract The complexity of forest structures plays a crucial role in regulating forest ecosystem functions and strongly influences biodiversity. Yet, knowledge of the global patterns and determinants of forest structural complexity remains scarce. Using a stand structural complexity index based on terrestrial laser scanning, we quantify the structural complexity of boreal, temperate, subtropical and tropical primary forests. We find that the global variation of forest structural complexity is largely explained by annual precipitation and precipitation seasonality (R² = 0.89). Using the structural complexity of primary forests as benchmark, we model the potential structural complexity across biomes and present a global map of the potential structural complexity of the earth´s forest ecoregions. Our analyses reveal distinct latitudinal patterns of forest structure and show that hotspots of high structural complexity coincide with hotspots of plant diversity. Considering the mechanistic underpinnings of forest structural complexity, our results suggest spatially contrasting changes of forest structure with climate change within and across biomes.more » « less
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