Life habits and evolutionary biology of new two-winged long-proboscid scorpionflies from mid-Cretaceous Myanmar amber

Long-proboscid scorpionflies are enigmatic, mid-Mesozoic insects associated with gymnosperm pollination. One major lineage, Aneuretopsychina, consists of four families plus two haustellate clades, Diptera and Siphonaptera. One clade, Pseudopolycentropodidae, from mid-Cretaceous Myanmar amber, contains Parapolycentropus. Here, we newly establish Dualula, assigned to Dualulidae, constituting the fifth lineage. Parapolycentropus and Dualula lineages are small, two-winged, with unique siphonate mouthparts for imbibing pollination drops. A cibarial pump provides siphonal food inflow; in Dualula, the siphon base surrounds a hypopharynx housing a small, valved pump constricted to a narrow salivary duct supplying outgoing enzymes for food fluidization. Indirect evidence links long-proboscid mouthpart structure with contemporaneous tubulate ovulate organs. Direct evidence of gymnospermous Cycadopites pollen is associated with one Parapolycentropus specimen. Parapolycentropus and Dualula exhibit hind-wing reduction that would precede haltere formation, likely caused by Ultrabithorax. Distinctive, male Aneuretopsychina genitalia are evident from specimens in copulo, supplemented by mixed-sex individuals of likely male mating swarms.