How did the Greek cities of Ionia come under Persian rule?

The Greek cities of Ionia came under Persian rule following the collapse of the Lydian Empire. Prior to the Persian conquest, most of these cities had been brought under the control of the Lydian king Croesus.1

When Cyrus the Great defeated Croesus and captured the Lydian capital of Sardis around 546 BCE, the Ionian Greek cities—formerly vassals of the Lydian king—became subject to the Achaemenid Empire. While Miletus had initially sided with the Persians during the conflict with Lydia, many other Greek towns were brought under Persian control through force, often led by the Median general Harpagus, who conducted the conquest of western Asia Minor on behalf of Cyrus.2345

Following their subjugation, these cities generally maintained a semi-autonomous status while recognizing Persian overlordship, though they were required to pay tribute and provide military service, including ships and troops for the Persian navy.43