The largest insects to have ever lived were the giant meganeurids of the Late Palaeozoic, ancient stem relatives of our modern dragonflies. With wingspans up to 71 cm, these iconic insects have been the subject of varied documentaries on Palaeozoic life, depicting them as patrolling for prey through coal swamp forests amid giant lycopsids, and cordaites. Such reconstructions are speculative as few definitive details of giant dragonfly biology are known. Most specimens of giant dragonflies are known from wings or isolated elements, but Meganeurites gracilipes preserves critical body structures, most notably those of the head. Here we show that it is unlikely it thrived in densely forested environments where its elongate wings would have become easily damaged. Instead, the species lived in more open habitats and possessed greatly enlarged compound eyes. These were dorsally hypertrophied, a specialization for long-distance vision above the animal in flight, a trait convergent with modern hawker dragonflies. Sturdy mandibles with acute teeth, strong spines on tibiae and tarsi, and a pronounced thoracic skewness are identical to those specializations used by dragonflies in capturing prey while in flight. The Palaeozoic Odonatoptera thus exhibited considerable morphological specializations associated with behaviours attributable to 'hawkers' or 'perchers' among extant Odonata.

Department of Zoology Faculty of Science Charles University Viničná 7 CZ 128 00 Praha 2 Czech Republic

Division of Entomology Natural History Museum and Department of Ecology and Evolutionary Biology 1501 Crestline Drive Suite 140 University of Kansas Lawrence Kansas 66045 4415 USA Division of Invertebrate Zoology American Museum of Natural History Central Park West at 79th Street New York New York 10024 5192 USA

Institut de Systématique Evolution Biodiversité ISYEB UMR 7205 CNRS MNHN UPMC EPHE Muséum national d'Histoire naturelle Sorbonne Universités 57 rue Cuvier CP 50 Entomologie F 75005 Paris France

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