Animals that glide produce aerodynamic forces that enable transit through the air in both arboreal and aquatic environments. The relative ease of gliding compared with flapping flight has led to a large diversity of taxa that have evolved some degree of flight capability. Glide paths are curved, reflecting the changing forces on the animal as it progresses through its aerial trajectory. These changing forces can be under control of the glider, which uses specific aspects of anatomy to modulate lift, drag, and rotational moments on the body. However, gliders share no single anatomical or behavioral feature, and some species are unspecialized for gliding, producing aerodynamic forces using posture and orientation alone. Animals use gliding in a broad range of ecological roles, suggesting that multiple performance metrics are relevant for consideration, but we are only beginning to understand how gliders produce and control their flight from takeoff to landing. In this review, we focus on the physical aspects of how glide trajectories are produced, and additionally discuss the range of morphologies and postures that are used to control aerial movements across the broad diversity of animal gliders.
In contrast to the patagial membranes of gliding vertebrates, the aerodynamic surfaces used by falling wingless ants to direct their aerial descent are unknown. We conducted ablation experiments to assess the relative contributions of the hindlegs, midlegs and gaster to gliding success in workers of the Neotropical arboreal ant Cephalotes atratus (Hymenoptera: Formicidae). Removal of hindlegs significantly reduced the success rate of directed aerial descent as well as the glide index for successful flights. Removal of the gaster alone did not significantly alter performance relative to controls. Equilibrium glide angles during successful targeting to vertical columns were statistically equivalent between control ants and ants with either the gaster or the hindlegs removed. High-speed video recordings suggested possible use of bilaterally asymmetric motions of the hindlegs to effect body rotations about the vertical axis during targeting manoeuvre. Overall, the control of gliding flight was remarkably robust to dramatic anatomical perturbations, suggesting effective control mechanisms in the face of adverse initial conditions (e.g. falling upside down), variable targeting decisions and turbulent wind gusts during flight.
Directed aerial descent (DAD) is used by a variety of arboreal animals to escape predators, to remain in the canopy, and to access resources. Here, we build upon the discovery of DAD in ants of tropical canopies by summarizing its known phylogenetic distribution among ant genera, and within both the subfamily Pseudomyrmecinae and the genus Cephalotes. DAD has multiple evolutionary origins in ants, occurring independently in numerous genera in the subfamilies Myrmicinae, Formicinae, and Pseudomyrmecinae. Ablation experiments and video recordings of ants in a vertical wind tunnel showed that DAD in Cephalotes atratus is achieved via postural changes, specifically orientation of the legs and gaster. The occurrence of DAD in Formicinae indicates that the presence of a postpetiole is not essential for the behavior. Evidence to date indicates that gliding behavior is accomplished by visual targeting mediated by the compound eyes, and is restricted to diurnally active ants that nest in trees. Occlusion of ocelli in Pseudomyrmex gracilis workers had no effect on their success or performance in gliding. Experimental assessment of the fate of ants that fall to the understory showed that ants landing in water are 15 times more likely to suffer lethal attacks than are ants landing in leaf litter. Variation in both the aerodynamic mechanisms and selective advantages of DAD merits further study given the broad taxonomic diversity of arboreal ants that engage in this intriguing form of flight.
The behaviour of directed aerial descent has been described for numerous taxa of wingless hexapods as they fall from the tropical rainforest canopy, but is not known in other terrestrial arthropods. Here, we describe similar controlled aerial behaviours for large arboreal spiders in the genus Selenops (Selenopidae). We dropped 59 such spiders from either canopy platforms or tree crowns in Panama and Peru; the majority (93%) directed their aerial trajectories towards and then landed upon nearby tree trunks. Following initial dorsoventral righting when necessary, falling spiders oriented themselves and then translated head-first towards targets; directional changes were correlated with bilaterally asymmetric motions of the anterolaterally extended forelegs. Aerial performance (i.e. the glide index) decreased with increasing body mass and wing loading, but not with projected surface area of the spider. Along with the occurrence of directed aerial descent in ants, jumping bristletails, and other wingless hexapods, this discovery of targeted gliding in selenopid spiders further indicates strong selective pressures against uncontrolled falls into the understory for arboreal taxa.
Gliding ants avoid predatory attacks and potentially mortal consequences of dislodgement from rainforest canopy substrates by directing their aerial descent towards nearby tree trunks. The ecologically relevant measure of performance for gliding ants is the ratio of net horizontal to vertical distance traveled over the course of a gliding trajectory, or glide index. To study variation in glide index, we measured three-dimensional trajectories of Cephalotes atratus ants gliding in natural rainforest habitats. We determined that righting phase duration, glide angle, and path directness all significantly influence variation in glide index. Unsuccessful landing attempts result in the ant bouncing off its target and being forced to make a second landing attempt. Our results indicate that ants are not passive gliders and that they exert active control over the aerodynamic forces they experience during their descent, despite their apparent lack of specialized control surfaces.
We develop a method to learn a bio-inspired motion control policy using data collected from hawkmoths navigating in a virtual forest. A Markov Decision Process (MDP) framework is introduced to model the dynamics of moths and sparse logistic regression is used to learn control policy parameters from the data. The results show that moths do not favor detailed obstacle location information in navigation, but rely heavily on optical flow. Using the policy learned from the moth data as a starting point, we propose an actor-critic learning algorithm to refine policy parameters and obtain a policy that can be used by an autonomous aerial vehicle operating in a cluttered environment. Compared with the moths' policy, the policy we obtain integrates both obstacle location and optical flow. We compare the performance of these two policies in terms of their ability to navigate in artificial forest areas. While the optimized policy can adjust its parameters to outperform the moth's policy in each different terrain, the moth's policy exhibits a high level of robustness across terrains.
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