Flight speed is positively correlated with body size in animals1. However, miniature featherwing beetles can fly at speeds and accelerations of insects three times their size2. Here we show that this performance results from a reduced wing mass and a previously unknown type of wing-motion cycle. Our experiment combines three-dimensional reconstructions of morphology and kinematics in one of the smallest insects, the beetle Paratuposa placentis (body length 395 μm). The flapping bristled wings follow a pronounced figure-of-eight loop that consists of subperpendicular up and down strokes followed by claps at stroke reversals above and below the body. The elytra act as inertial brakes that prevent excessive body oscillation. Computational analyses suggest functional decomposition of the wingbeat cycle into two power half strokes, which produce a large upward force, and two down-dragging recovery half strokes. In contrast to heavier membranous wings, the motion of bristled wings of the same size requires little inertial power. Muscle mechanical power requirements thus remain positive throughout the wingbeat cycle, making elastic energy storage obsolete. These adaptations help to explain how extremely small insects have preserved good aerial performance during miniaturization, one of the factors of their evolutionary success.

Aerodynamic force generation capacity of the wing of a miniature beetle Paratuposa placentis is evaluated using a combined experimental and numerical approach. The wing has a peculiar shape reminiscent of a bird feather, often found in the smallest insects. Aerodynamic force coefficients are determined from a dynamically scaled force measurement experiment with rotating bristled and membrane wing models in a glycerin tank. Subsequently, they are used as numerical validation data for computational fluid dynamics simulations using an adaptive Navier–Stokes solver. The latter provides access to important flow properties such as leakiness and permeability. It is found that, in the considered biologically relevant regimes, the bristled wing functions as a less than $$50\%$$ 50 % leaky paddle, and it produces between 66 and $$96\%$$ 96 % of the aerodynamic drag force of an equivalent membrane wing. The discrepancy increases with increasing Reynolds number. It is shown that about half of the aerodynamic normal force exerted on a bristled wing is due to viscous shear stress. The paddling effectiveness factor is proposed as a measure of aerodynamic efficiency. Graphic abstract

Size is a key to locomotion. In insects, miniaturization leads to fundamental changes in wing structure and kinematics, making the study of flight in the smallest species important for basic biology and physics, and, potentially, for applied disciplines. However, the flight efficiency of miniature insects has never been studied, and their speed and maneuverability have remained unknown. We report a comparative study of speeds and accelerations in the smallest free-living insects, featherwing beetles (Coleoptera: Ptiliidae), and in larger representatives of related groups of Staphylinoidea. Our results show that the average and maximum flight speeds of larger ptiliids are extraordinarily high and comparable to those of staphylinids that have bodies 3 times as long. This is one of the few known exceptions to the “Great Flight Diagram,” according to which the flight speed of smaller organisms is generally lower than that of larger ones. The horizontal acceleration values recorded in Ptiliidae are almost twice as high as even in Silphidae, which are more than an order of magnitude larger. High absolute and record-breaking relative flight characteristics suggest that the unique morphology and kinematics of the ptiliid wings are effective adaptations to flight at low Reynolds numbers. These results are important for understanding the evolution of body size and flight in insects and pose a challenge to designers of miniature biomorphic aircraft.

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The diagnoses of Liadytidae Ponomarenko, 1977, Liadytiscinae Prokin & Ren, 2010, Liadytiscus Prokin & Ren, 2010 and Mesoderus Prokin & Ren, 2010 (Dytiscidae) are modified, and the following new taxa are described from Mesozoic fossils: Liadytes aspidytoides sp. n. (Liadytidae); Mesoderini trib. n., Liadyxianus kirejtshuki gen. n. et sp. n., Mesoderus punctatus sp. n., Mesoderus ovatus sp. n., Mesodytes rhantoides gen. n. et sp. n., Palaeodytes baissiensis sp. n. and Cretodytes incertus sp. n. (Dytiscidae). A summarized checklist of all Mesozoic Liadytidae and Dytiscidae known from adults is given, and an identification key to the genera of Mesozoic Dytiscidae known from adults is provided for the first time. Palaeodytes incompletus Ponomarenko, Coram & Jarzembowski, 2005 (the suffix of the specific epithet is emended from the original incompleta) is found to belong not to this genus, but to another one, which remains to be described. The fossil larva Angaragabus jurassicus Ponomarenko, 1963 from the Lower Jurassic of Irkutsk Oblast, Russia, probably belonging to Liadytidae, is re-examined. If this larva actually belongs to Liadytidae, then its morphological characters provide additional confirmation of the conclusion, based on the characters of adult liadytids, that the family is quite separate from the recent family Aspidytidae, and the similarity between the adults of both families results from parallel processes in the evolution of the superfamily Dytiscoidea. We show that the principal trends of morphological changes of Liadytidae and Dytiscidae during the Upper Jurassic and Lower Cretaceous included a consistent increase in the area of the metacoxal plates at the expense of decreasing area of the lateral lobes of the metaventrite ("wings"), flattening and loss of the lateral border of the elevated median area of the metaventrite, and shortening and dilation of the metafemur and metatibia. These changes were probably associated with an increased load of swimming taken by the hindlegs, which required, among other things, the development of swimming muscles attached to the metacoxal plates. The development of the hindlegs allowed diving deeper, thus being an adaptation to the nektonic instead of benthic lifestyle. This is confirmed by the adaptive coloration of the Liadytidae and Dytiscidae found in Shar Teg (Liadytes aspidytoides sp. n.), Yixian (Mesoderus magnus Prokin & Ren, 2010) and Baisa (Palaeodytes baissiensis sp. n.), in which the dorsum was darker than underside, providing camouflage in the depths of the water.