Extant cicada (Hemiptera: Cicadoidea) includes widely distributed Cicadidae and relictual Tettigarctidae, with fossils ascribed to these two groups based on several distinct, minimally varying morphological differences that define their extant counterparts. However, directly assigning Mesozoic fossils to modern taxa may overlook the role of unique and transitional features provided by fossils in tracking their early evolutionary paths. Here, based on adult and nymphal fossils from mid-Cretaceous Kachin amber of Myanmar, we explore the phylogenetic relationships and morphological disparities of fossil and extant cicadoids. Our results suggest that Cicadidae and Tettigarctidae might have diverged at or by the Middle Jurassic, with morphological evolution possibly shaped by host plant changes. The discovery of tymbal structures and anatomical analysis of adult fossils indicate that mid-Cretaceous cicadas were silent as modern Tettigarctidae or could have produced faint tymbal-related sounds. The discovery of final-instar nymphal and exuviae cicadoid fossils with fossorial forelegs and piercing-sucking mouthparts indicates that they had most likely adopted a subterranean lifestyle by the mid-Cretaceous, occupying the ecological niche of underground feeding on root. Our study traces the morphological, behavioral, and ecological evolution of Cicadoidea from the Mesozoic, emphasizing their adaptive traits and interactions with their living environments.

Australian Museum Research Institute Sydney NSW 2010 Australia

Beijing Xiachong Amber Museum Beijing 100083 China

Department of Entomology and Behavior Ecology Evolution and Systematics Program University of Maryland College Park MD 20742 USA

Department of Geological Survey and Mineral Exploration Ministry of Natural Resources and Environmental Conservation Myanmar Gems Museum Nay Pyi Taw 15011 Myanmar

Department of Paleobiology National Museum of Natural History Smithsonian Institution Washington DC 20013 USA

Institute of Geology and Paleontology Charles University Prague 12843 Czech Republic

Institute of Geology and Paleontology Linyi University Linyi 276000 China

Key Laboratory of Plant Protection Resources and Pest Management of the Ministry of Education College of Plant Protection Northwest A and F University Yangling Shaanxi 712100 China

Laboratory of Evolutionary Entomology and Museum of Amber Inclusions Department of Invertebrate Zoology and Parasitology University of Gdańsk Gdańsk PL80 308 Poland

Princeton Consultants Princeton NJ 08540 USA

School of Life Sciences Capital Normal University Beijing 100048 China

Section Palaeontology Institute of Geosciences Rheinische Friedrich Wilhelms Universität Bonn Bonn 53115 Germany

State Key Laboratory of Paleobiology and Stratigraphy Nanjing Institute of Geology and Palaeontology and Center for Excellence in Life and Paleoenvironment Chinese Academy of Sciences Nanjing 210008 China

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