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In 499 B.C.E. - the Ionian Greeks in Anatolia - who had been invaded by the Persians in 546 B.C.E. - rebelled against Persian control and were aided by the city-state ___ on the Greek mainland.

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1. In 499 B.C.E. - the Ionian Greeks in Anatolia - who had been invaded by the Persians in 546 B.C.E. - rebelled against Persian control and were aided by the city-state ___ on the Greek mainland.





 
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