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1. Adaptive shifts underlie the divergence in wing morphology in bombycoid moths

   https://doi.org/10.1098/rspb.2021.0677  

   Aiello, Brett R. ; Tan, Milton ; Bin Sikandar, Usama ; Alvey, Alexis J. ; Bhinderwala, Burhanuddin ; Kimball, Katalina C. ; Barber, Jesse R. ; Hamilton, Chris A. ; Kawahara, Akito Y. ; Sponberg, Simon ( August 2021 , Proceedings of the Royal Society B: Biological Sciences)

   null (Ed.)

   The evolution of flapping flight is linked to the prolific success of insects. Across Insecta, wing morphology diversified, strongly impacting aerodynamic performance. In the presence of ecological opportunity, discrete adaptive shifts and early bursts are two processes hypothesized to give rise to exceptional morphological diversification. Here, we use the sister-families Sphingidae and Saturniidae to answer how the evolution of aerodynamically important traits is linked to clade divergence and through what process(es) these traits evolve. Many agile Sphingidae evolved hover feeding behaviours, while adult Saturniidae lack functional mouth parts and rely on a fixed energy budget as adults. We find that Sphingidae underwent an adaptive shift in wing morphology coincident with life history and behaviour divergence, evolving small high aspect ratio wings advantageous for power reduction that can be moved at high frequencies, beneficial for flight control. By contrast, Saturniidae, which do not feed as adults, evolved large wings and morphology which surprisingly does not reduce aerodynamic power, but could contribute to their erratic flight behaviour, aiding in predator avoidance. We suggest that after the evolution of flapping flight, diversification of wing morphology can be potentiated by adaptative shifts, shaping the diversity of wing morphology across insects. 

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2. Effects of wing pitch kinematics on both aerodynamic and olfactory functions in an upwind surge

   https://doi.org/10.1177/0954406220907950  

   Li, Chengyu ( January 2021 , Proceedings of the Institution of Mechanical Engineers, Part C: Journal of Mechanical Engineering Science)

   Flapping wings of insects serve for both generating aerodynamic forces and enhancing olfactory sensitivities when navigating on the odor-rich planet. Despite the extensive investigations of the aerodynamic function of flapping wings, we have limited understanding of how the flapping wings potentially affect the physiological sensitivities during flight. In this paper, direct numerical simulations were used to investigate a fruit fly model in an upwind surging motion. The wing pitch kinematics were prescribed using a hyperbolic function, which can change the wing pitch profile from a sinusoidal function to a step function by adjusting the “C” factor in the hyperbolic function. Both aerodynamic performance and olfactory detections were quantified at various wing pitch kinematics patterns. The effects of flapping wings on the odor transport were visualized using the Lagrangian approach by uniformly releasing passive odor tracers in upstream. The study revealed that the insect had the potential to achieve higher aerodynamic performance by tailoring wing pitch kinematics, but it could reduce the odor mass flux around the antenna. It was suspected that the natural flyers might sacrifice certain aerodynamic potential to enhance their olfactory sensitivity for surviving purposes. In addition, a trap-and-flick mechanism is proposed here during the supination phase in order to enhance the olfactory sensitivity. Similar to the clip-and-fling mechanism for enhancing the force generation during the pronation phase, the newly proposed trap-and-flick mechanism is also due to the wing-wing interaction in flapping flight. These findings could provide important implications for engineering applications of odor-guided flapping flight.

    

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3. The hawkmoth wingbeat is not at resonance

   https://doi.org/10.1098/rsbl.2022.0063  

   Gau, Jeff ; Wold, Ethan S. ; Lynch, James ; Gravish, Nick ; Sponberg, Simon ( May 2022 , Biology Letters)

   Flying insects have elastic materials within their exoskeletons that could reduce the energetic cost of flight if their wingbeat frequency is matched to a mechanical resonance frequency. Flapping at resonance may be essential across flying insects because of the power demands of small-scale flapping flight. However, building up large-amplitude resonant wingbeats over many wingstrokes may be detrimental for control if the total mechanical energy in the spring-wing system exceeds the per-cycle work capacity of the flight musculature. While the mechanics of the insect flight apparatus can behave as a resonant system, the question of whether insects flap their wings at their resonant frequency remains unanswered. Using previous measurements of body stiffness in the hawkmoth, Manduca sexta , we develop a mechanical model of spring-wing resonance with aerodynamic damping and characterize the hawkmoth's resonant frequency. We find that the hawkmoth's wingbeat frequency is approximately 80% above resonance and remains so when accounting for uncertainty in model parameters. In this regime, hawkmoths may still benefit from elastic energy exchange while enabling control of aerodynamic forces via frequency modulation. We conclude that, while insects use resonant mechanics, tuning wingbeats to a simple resonance peak is not a necessary feature for all centimetre-scale flapping flyers. 

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4. Experimental studies suggest differences in the distribution of thorax elasticity between insects with synchronous and asynchronous musculature

   https://doi.org/10.1098/rsif.2023.0029  

   Casey, Cailin ; Heveran, Chelsea ; Jankauski, Mark ( April 2023 , Journal of The Royal Society Interface)

   Insects have developed diverse flight actuation mechanisms, including synchronous and asynchronous musculature. Indirect actuation, used by insects with both synchronous and asynchronous musculature, transforms thorax exoskeletal deformation into wing rotation. Though thorax deformation is often attributed exclusively to muscle tension, the inertial and aerodynamic forces generated by the flapping wings may also contribute. In this study, a tethered flight experiment was used to simultaneously measure thorax deformation and the inertial/aerodynamic forces acting on the thorax generated by the flapping wing. Compared to insects with synchronous musculature, insects with asynchronous muscle deformed their thorax 60% less relative to their thorax diameter and their wings generated 2.8 times greater forces relative to their body weight. In a second experiment, dorsalventral thorax stiffness was measured across species. Accounting for weight and size, the asynchronous thorax was on average 3.8 times stiffer than the synchronous thorax in the dorsalventral direction. Differences in thorax stiffness and forces acting at the wing hinge led us to hypothesize about differing roles of series and parallel elasticity in the thoraxes of insects with synchronous and asynchronous musculature. Specifically, wing hinge elasticity may contribute more to wing motion in insects with asynchronous musculature than in those with synchronous musculature. 

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5. Haemolymph viscosity in hawkmoths and its implications for hovering flight

   https://doi.org/10.1098/rspb.2022.2185  

   Brasovs, Artis ; Palaoro, Alexandre V. ; Aprelev, Pavel ; Beard, Charles E. ; Adler, Peter H. ; Kornev, Konstantin G. ( April 2023 , Proceedings of the Royal Society B: Biological Sciences)

   Viscosity determines the resistance of haemolymph flow through the insect body. For flying insects, viscosity is a major physiological parameter limiting flight performance by controlling the flow rate of fuel to the flight muscles, circulating nutrients and rapidly removing metabolic waste products. The more viscous the haemolymph, the greater the metabolic energy needed to pump it through confined spaces. By employing magnetic rotational spectroscopy with nickel nanorods, we showed that viscosity of haemolymph in resting hawkmoths (Sphingidae) depends on wing size non-monotonically. Viscosity increases for small hawkmoths with high wingbeat frequencies, reaches a maximum for middle-sized hawkmoths with moderate wingbeat frequencies, and decreases in large hawkmoths with slower wingbeat frequencies but greater lift. Accordingly, hawkmoths with small and large wings have viscosities approaching that of water, whereas hawkmoths with mid-sized wings have more than twofold greater viscosity. The metabolic demands of flight correlate with significant changes in circulatory strategies via modulation of haemolymph viscosity. Thus, the evolution of hovering flight would require fine-tuned viscosity adjustments to balance the need for the haemolymph to carry more fuel to the flight muscles while decreasing the viscous dissipation associated with its circulation. 

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