Exceptional preservation and foot structure reveal ecological transitions and lifestyles of early theropod flyers

Morphology of keratinised toe pads and foot scales, hinging of foot joints and claw shape and size all inform the grasping ability, cursoriality and feeding mode of living birds. Presented here is morphological evidence from the fossil feet of early theropod flyers. Foot soft tissues and joint articulations are qualitatively assessed using laser-stimulated fluorescence. Pedal claw shape and size are quantitatively analysed using traditional morphometrics. We interpret these foot data among existing evidence to better understand the evolutionary ecology of early theropod flyers. Jurassic flyers like Anchiornis and Archaeopteryx show adaptations suggestive of relatively ground-dwelling lifestyles. Early Cretaceous flyers then diversify into more aerial lifestyles, including generalists like Confuciusornis and specialists like the climbing Fortunguavis . Some early birds, like the Late Jurassic Berlin Archaeopteryx and Early Cretaceous Sapeornis , show complex ecologies seemingly unique among sampled modern birds. As a non-bird flyer, finding affinities of Microraptor to a more specialised raptorial lifestyle is unexpected. Its hawk-like characteristics are rare among known theropod flyers of the time suggesting that some non-bird flyers perform specialised roles filled by birds today. We demonstrate diverse ecological profiles among early theropod flyers, changing as flight developed, and some non-bird flyers have more complex ecological roles. The shape of bird toe pads and foot scales can be used to infer their behaviour. Here, the authors examine fossil evidence of toe pads and scales, in addition to claws and bones, from birds and close relatives, illustrating diverse lifestyles and ecological roles among early theropod flyers.