The Late Palaeozoic insect superorder Palaeodictyopterida exhibits a remarkable disparity of larval ecomorphotypes, enabling these animals to occupy diverse ecological niches. The widely accepted hypothesis presumed that their immature stages only occupied terrestrial habitats, although authors more than a century ago hypothesized they had specializations for amphibious or even aquatic life histories. Here, we show that different species had a disparity of semiaquatic or aquatic specializations in larvae and even the supposed retention of abdominal tracheal gills by some adults. While a majority of mature larvae in Palaeodictyoptera lack unambiguous lateral tracheal gills, some recently discovered early instars had terminal appendages with prominent lateral lamellae like in living damselflies, allowing support in locomotion along with respiratory function. These results demonstrate that some species of Palaeodictyopterida had aquatic or semiaquatic larvae during at least a brief period of their post-embryonic development. The retention of functional gills or gill sockets by adults indicates their amphibious lifestyle and habitats tightly connected with a water environment as is analogously known for some modern Ephemeroptera or Plecoptera. Our study refutes an entirely terrestrial lifestyle for all representatives of the early diverging pterygote group of Palaeodictyopterida, a greatly varied and diverse lineage which probably encompassed many different biologies and life histories.

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 University of Kansas 1501 Crestline Drive Suite 140 Lawrence KS 66045 USA

Division of Invertebrate Zoology American Museum of Natural History Central Park West at 79th Street New York NY 10024 5192 USA

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

Institute of Systematics and Evolution of Animals Polish Academy of Sciences ul Sławkowska 17 31 016 Kraków Poland

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