Fleas are one of the major lineages of ectoparasitic insects and are now highly specialized for feeding on the blood of birds or mammals1. This has isolated them among holometabolan insect orders, although they derive from the Antliophora (scorpionflies and true flies). Like most ectoparasitic lineages, their fossil record is meagre and confined to Cenozoic-era representatives of modern families1, so that we lack evidence of the origins of fleas in the Mesozoic era. The origins of the first recognized Cretaceous stem-group flea,Tarwinia, remains highly controversial1. Here we report fossils of the oldest definitive fleas—giant forms from the Middle Jurassic and Early Cretaceous periods of China. They exhibit many defining features of fleas but retain primitive traits such as non-jumping hindlegs. More importantly, all have stout and elongate sucking siphons for piercing the hides of their hosts, implying that these fleas may be rooted among the pollinating ‘long siphonate’ scorpionflies of the Mesozoic. Their special morphology suggests that their earliest hosts were hairy or feathered ‘reptilians’,and that they radiated to mammalian and bird hosts later in the Cenozoic.