Ionian Revolt The Ionian Revolt , and associated revolts in Aeolis , Doris , Cyprus and Caria , were military rebellions by several regions of Asia Minor against Persian rule , lasting from 499 BC to 493 BC . At the heart of the rebellion was the dissatisfaction of the Greek cities of Asia Minor with the tyrants appointed by Persia to rule them , along with the individual actions of two Milesian tyrants , Histiaeus and Aristagoras . The cities of Ionia had been conquered by Persia around 540 BC , and thereafter were ruled by native tyrants , nominated by the Persian satrap in Sardis . In 499 BC , the then tyrant of Miletus , Aristagoras , launched a joint expedition with the Persian satrap Artaphernes to conquer Naxos , in an attempt to bolster his position . The mission was a debacle , and sensing his imminent removal as tyrant , Aristagoras chose to incite the whole of Ionia into rebellion against the Persian king Darius the Great . In 498 BC , supported by troops from Athens and Eretria , the Ionians marched on , captured , and burnt Sardis . However , on their return journey to Ionia , they were followed by Persian troops , and decisively beaten at the Battle of Ephesus . This campaign was the only offensive action by the Ionians , who subsequently went on the defensive . The Persians responded in 497 BC with a three pronged attack aimed at recapturing the outlying areas of the rebellion , but the spread of the revolt to Caria meant that the largest army , under Daurises , relocated there . While initially campaigning successfully in Caria , this army was annihilated in an ambush at the Battle of Pedasus . This resulted in a stalemate for the rest of 496 BC and 495 BC . By 494 BC the Persian army and navy had regrouped , and they made straight for the epicentre of the rebellion at Miletus . The Ionian fleet sought to defend Miletus by sea , but were decisively beaten at the Battle of Lade , after the defection of the Samians . Miletus was then besieged , captured , and its population was brought under Persian rule . This double defeat effectively ended the revolt , and the Carians surrendered to the Persians as a result . The Persians spent 493 BC reducing the cities along the west coast that still held out against them , before finally imposing a peace settlement on Ionia which was generally considered to be both just and fair . The Ionian Revolt constituted the first major conflict between Greece and the Persian Empire , and as such represents the first phase of the Greco - Persian Wars . Although Asia Minor had been brought back into the Persian fold , Darius vowed to punish Athens and Eretria for their support of the revolt . Moreover , seeing that the myriad city states of Greece posed a continued threat to the stability of his Empire , according to Herodotus , Darius decided to conquer the whole of Greece . In 492 BC , the first Persian invasion of Greece , the next phase of the Greco - Persian Wars , would begin as a direct consequence of the Ionian Revolt . Sources . Practically the only primary source for the Ionian Revolt is the Greek historian Herodotus . Herodotus , who has been called the ' Father of History ' , was born in 484 BC in Halicarnassus , Asia Minor ( then under Persian overlordship ) . He wrote his ' Enquiries ' ( Greek — " Historia " ; English — " ( The ) Histories " ) around 440 – 430 BC , trying to trace the origins of the Greco - Persian Wars , which would still have been relatively recent history ( the wars finally ending in 450 BC ) . Herodotus ' s approach was entirely novel , and at least from the point of view of Western society , he does seem to have invented ' history ' as we know it . As Holland has it : " For the first time , a chronicler set himself to trace the origins of a conflict not to a past so remote so as to be utterly fabulous , nor to the whims and wishes of some god , nor to a people ' s claim to manifest destiny , but rather explanations he could verify personally . " Some subsequent ancient historians , despite following in his footsteps , criticised Herodotus , starting with Thucydides . Nevertheless , Thucydides chose to begin his history where Herodotus left off ( at the Siege of Sestos ) , and therefore presumably felt that Herodotus ' s history was accurate enough not to need re - writing or correcting . Plutarch criticised Herodotus in his essay " On The Malignity of Herodotus " , describing Herodotus as " " Philobarbaros " " ( barbarian - lover ) and for not being pro - Greek enough , which suggests that Herodotus might actually have done a reasonable job of being even - handed . A negative view of Herodotus was passed on to Renaissance Europe , though he remained widely read . However , since the 19th century his reputation has been dramatically rehabilitated by the age of democracy and some archaeological finds which have repeatedly confirmed his version of events . The prevailing modern view is that Herodotus generally did a remarkable job in his " Historia " , but that some of his specific details ( particularly troop numbers and dates ) should be viewed with skepticism . Nevertheless , there are still many historians who believe Herodotus ' account has an anti - Persian bias and that much of his story was embellished for dramatic effect . Background . In the dark age that followed the collapse of the Mycenaean civilization , significant numbers of Greeks emigrated to Asia Minor and settled there . These settlers were from three tribal groups : the Aeolians , Dorians and Ionians . The Ionians had settled along the coasts of Lydia and Caria , founding the twelve cities which made up Ionia . These cities were Miletus , Myus and Priene in Caria ; Ephesus , Colophon , Lebedos , Teos , Clazomenae , Phocaea and Erythrae in Lydia ; and the islands of Samos and Chios . Although the Ionian cities were independent from each other , they acknowledged their shared heritage , and had a common temple and meeting place , the Panionion . They thus formed a ' cultural league ' , to which they would admit no other cities , or even other tribal Ionians . The cities of Ionia had remained independent until they were conquered by the famous Lydian king Croesus , in around 560 BC . The Ionian cities then remained under Lydian rule until Lydia was in turn conquered by the nascent Achaemenid Empire of Cyrus the Great . While fighting the Lydians , Cyrus had sent messages to the Ionians asking them to revolt against Lydian rule , which the Ionians had refused to do . After Cyrus completed the conquest of Lydia , the Ionian cities now offered to be his subjects under the same terms as they had been subjects of Croesus . Cyrus refused , citing the Ionians ' unwillingness to help him previously . The Ionians thus prepared to defend themselves , and Cyrus sent the Median general Harpagus to conquer Ionia . He first attacked Phocaea ; the Phocaeans decided to entirely abandon their city and sail into exile in Sicily , rather than become Persian subjects ( although many subsequently returned ) . Some Teians also chose to emigrate when Harpagus attacked Teos , but the rest of the Ionians remained , and were in turn conquered . The Persians found the Ionians difficult to rule . Elsewhere in the empire , Cyrus was able to identify elite native groups to help him rule his new subjects – such as the priesthood of Judea . No such group existed in Greek cities at this time ; while there was usually an aristocracy , this was inevitably divided into feuding factions . The Persians thus settled for sponsoring a tyrant in each Ionian city , even though this drew them into the Ionians ' internal conflicts . Furthermore , a tyrant might develop an independent streak , and have to be replaced . The tyrants themselves faced a difficult task ; they had to deflect the worst of their fellow citizens ' hatred , while staying in the favour of the Persians . About 40 years after the Persian conquest of Ionia , and in the reign of the fourth Persian king , Darius the Great , the stand - in Milesian tyrant Aristagoras found himself in this familiar predicament . Aristagoras ' s uncle Histiaeus had accompanied Darius on campaign in 513 BC , and when offered a reward , had asked for part of the conquered Thracian territory . Although this was granted , Histiaeus ' s ambition alarmed Darius ' s advisors , and Histiaeus was thus further ' rewarded ' by being compelled to remain in Susa as Darius ' s " Royal Table - Companion " . Taking over from Histiaeus , Aristagoras was faced with bubbling discontent in Miletus . In 500 BC , Aristagoras was approached by some exiles from Naxos , who asked him to take control of the island . Seeing an opportunity to strengthen his position in Miletus by conquering Naxos , Aristagoras approached the satrap of Lydia , Artaphernes , with a proposal . If Artaphernes provided an army , Aristagoras would conquer the island , thus extending the boundaries of the empire for Darius , and he would then give Artaphernes a share of the spoils to cover the cost of raising the army . Artaphernes agreed in principle , and asked Darius for permission to launch the expedition . Darius assented to this , and a force of 200 triremes was assembled in order to attack Naxos the following year . Naxos campaign ( 499 BC ) . In the spring of 499 BC , Artaphernes readied the Persian force , and placed his cousin Megabates in command . He then sent ships on to Miletus , where the Ionian troops levied by Aristagoras embarked , and the force then set sail for Naxos . The expedition quickly descended into a debacle . Aristagoras fell out with Megabates on the journey towards Naxos , and Herodotus says that Megabates then sent messengers to Naxos , warning the Naxians of the forces ' s intention . It is also possible , however , that this story was spread by Aristagoras after the event , by way of a justification for the subsequent failure of the campaign . At any rate , the Naxians were able to prepare properly for a siege , and the Persians arrived to a well - defended expedition . The Persians laid siege to the Naxians for four months , but eventually they and Aristagoras both ran out of money . The force therefore sailed despondently back to the mainland . Start of the Ionian Revolt ( 499 BC ) . With the failure of his attempt to conquer Naxos , Aristagoras found himself in dire straits ; he was unable to repay Artaphernes , and had , moreover , alienated himself from the Persian royal family . He fully expected to be stripped of his position by Artaphernes . In a desperate attempt to save himself , Aristagoras chose to incite his own subjects , the Milesians , to revolt against their Persian masters , thereby beginning the Ionian Revolt . In autumn 499 BC , Aristagoras held a meeting with the members of his faction in Miletus . He declared that in his opinion the Milesians should revolt , to which all but the historian Hecataeus agreed . At the same time , a messenger sent by Histiaeus arrived in Miletus , imploring Aristagoras to rebel against Darius . Herodotus suggests that this was because Histiaeus was desperate to return to Ionia , and thought he would be sent to Ionia if there was a rebellion . Aristagoras therefore openly declared his revolt against Darius , abdicated from his role as tyrant , and declared Miletus to be a democracy . Herodotus has no doubt that this was only a pretence on Aristagoras ' s part of giving up power . Rather it was designed to make the Milesians enthusiastically join the rebellion . The army that had been sent to Naxos was still assembled at Myus and included contingents from other Greek cities of Asia Minor ( i . e . Aeolia and Doris ) as well as men from Mytilene , Mylasa , Temera and Cyme . Aristagoras sent men to capture all the Greek tyrants present in the army and handed them over to their respective cities in order to gain the cooperation of those cities . Some of the tyrants were executed by their cities , but most were simply banished . It has also been suggested ( Herodotus does not explicitly say so ) that Aristagoras incited the whole army to join his revolt , and also took possession of the ships that the Persians had supplied . If the latter is true , it may explain the length of time it took for the Persians to launch a naval assault on Ionia , since they would have needed to build a new fleet . Although Herodotus presents the revolt as a consequence of Aristagoras and Histiaeus ' s personal motives , it is clear that Ionia must have been ripe for rebellion anyway . The primary grievance was the tyrants installed by the Persians . While Greek states had in the past often been ruled by tyrants , this was a form of government on the decline . Moreover , past tyrants had tended ( and needed ) to be strong and able leaders , whereas the rulers appointed by the Persians were simply the representatives of the Persians . Backed by Persian military might , these tyrants did not need the support of the population , and could thus rule absolutely . Aristagoras ' s actions have thus been likened to tossing a flame into a kindling box ; they incited rebellion across Ionia , and tyrannies were everywhere abolished , and democracies established in their place . Aristagoras had brought all of Hellenic Asia Minor into revolt , but evidently realised that the Greeks would need other allies in order to successfully fight the Persians . In the winter of 499 BC , he first sailed to Sparta , the pre - eminent Greek state in matters of war . However , despite Aristagoras ' s entreaties , the Spartan king Cleomenes I turned down the offer to lead the Greeks against the Persians . Aristagoras therefore turned instead to Athens . Athens had recently become a democracy , overthrowing its own tyrant Hippias . In their fight to establish the democracy , the Athenians had asked the Persians for aid ( which was not in the end needed ) , in return for submitting to Persian overlordship . Some years later , Hippias had attempted to regain power in Athens , assisted by the Spartans . This attempt failed and Hippias fled to Artaphernes , and tried to persuade him to subjugate Athens . The Athenians dispatched ambassadors to Artaphernes to dissuade him from taking action , but Artaphernes merely instructed the Athenians to take Hippias back as tyrant . Needless to say , the Athenians had baulked at this , and resolved instead to be openly at war with Persia . Since they were already an enemy of Persia , Athens was already in a position to support the Ionian cities in their revolt . The fact that the Ionian democracies were inspired by the example of the Athenian democracy no doubt helped persuade the Athenians to support the Ionian Revolt , especially since the cities of Ionia were ( supposedly ) originally Athenian colonies . Aristagoras was also successful in persuading the city of Eretria to send assistance to the Ionians for reasons that are not completely clear . Possibly commercial reasons were a factor ; Eretria was a mercantile city , whose trade was threatened by Persian dominance of the Aegean . Herodotus suggests that the Eretrians supported the revolt in order to repay the support the Milesians had given Eretria some time previously , possibly referring to the Lelantine War . Ionian offensive ( 498 BC ) . Over the winter , Aristagoras continued to foment rebellion . In one incident , he told a group of Paeonians ( originally from Thrace ) , who Darius had brought to live in Phrygia , to return to their homeland . Herodotus says that his only purpose in doing this was to vex the Persian high command . Sardis . In the spring of 498 BC , an Athenian force of twenty triremes , accompanied by five from Eretria , set sail for Ionia . They joined up with the main Ionian force near Ephesus . Declining to personally lead the force , Aristagoras appointed his brother Charopinus and another Milesian , Hermophantus , as generals . This force was then guided by the Ephesians through the mountains to Sardis , Artaphernes ' s satrapal capital . The Greeks caught the Persians unaware , and were able to capture the lower city . However , Artaphernes still held the citadel with a significant force of men . The lower city then caught on fire , Herodotus suggests accidentally , which quickly spread . The Persians in the citadel , being surrounded by a burning city , emerged into the market - place of Sardis , where they fought with the Greeks , forcing them back . The Greeks , demoralised , then retreated from the city , and began to make their way back to Ephesus . Herodotus reports that when Darius heard of the burning of Sardis , he swore vengeance upon the Athenians ( after asking who they indeed were ) , and tasked a servant with reminding him three times each day of his vow : " Master , remember the Athenians " . Battle of Ephesus . Herodotus says that when the Persians in Asia Minor heard of the attack on Sardis , they gathered together , and marched to the relief of Artaphernes . When they arrived at Sardis , they found the Greeks recently departed . So they followed their tracks back towards Ephesus . They caught up with the Greeks outside Ephesus and the Greeks were forced to turn and prepare to fight . Holland suggests that the Persians were primarily cavalry ( hence their ability to catch up with the Greeks ) . The typical Persian cavalry of the time were probably missile cavalry , whose tactics were to wear down a static enemy with volley after volley . It is clear that the demoralised and tired Greeks were no match for the Persians , and were completely routed in the battle which ensued at Ephesus . Many were killed , including the Eretrian general , Eualcides . The Ionians who escaped the battle made for their own cities , while the remaining Athenians and Eretrians managed to return to their ships and sailed back to Greece . Spread of the revolt . The Athenians now ended their alliance with the Ionians , since the Persians had proved to be anything but the easy prey that Aristagoras had described . However , the Ionians remained committed to their rebellion and the Persians did not seem to follow up their victory at Ephesus . Presumably these " ad hoc " forces were not equipped to lay siege to any of the cities . Despite the defeat at Ephesus , the revolt actually spread further . The Ionians sent men to the Hellespont and Propontis and captured Byzantium and the other nearby cities . They also persuaded the Carians to join the rebellion . Furthermore , seeing the spread of the rebellion , the kingdoms of Cyprus also revolted against Persian rule without any outside persuasion . Persian counter - offensive ( 497 – 495 BC ) . Herodotus ' s narrative after the Battle of Ephesus is ambiguous in its exact chronology ; historians generally place Sardis and Ephesus in 498 BC . Herodotus next describes the spread of the revolt ( thus also in 498 BC ) , and says that the Cypriots had one year of freedom , therefore placing the action in Cyprus to 497 BC . He next says that " Daurises , Hymaees , and Otanes , all of them Persian generals and married to daughters of Darius , pursued those Ionians who had marched to Sardis , and drove them to their ships . After this victory they divided the cities among themselves and sacked them . " This passage implies these Persian generals counter - attacked " immediately " after the Battle of Ephesus . However , the cities that Herodotus describes Daurises as besieging were on the Hellespont , which ( by Herodotus ' s own reckoning ) did not become involved in the revolt until " after " Ephesus . It is therefore easiest to reconcile the account by assuming that Daurises , Hymaees , and Otanes waited until the next campaigning season ( i . e . 497 BC ) , before going on the counter - offensive . The Persian actions that Herodotus describes at the Hellespont and in Caria seem to be in the same year , and most commentators place them in 497 BC . Cyprus . In Cyprus , all the kingdoms had revolted except that of Amathus . The leader of the Cypriot revolt was Onesilus , brother of the king of Salamis - in - Cyprus , Gorgus . Gorgus did not want to revolt , so Onesilus locked his brother out of the city and made himself king . Gorgus went over to the Persians , and Onesilus persuaded the other Cypriots , apart from the Amathusians , to revolt . He then settled down to besiege Amathus . The following year ( 497 BC ) , Onesilus ( still besieging Amathus ) , heard that a Persian force under Artybius had been dispatched to Cyprus . Onesilus thus sent messengers to Ionia , asking them to send reinforcements , which they did , " in great force " . A Persian army eventually arrived in Cyprus , supported by a Phonecia n fleet . The Ionians opted to fight at sea and defeated the Phoenicians . In the simultaneous land battle , the Cypriots gained an initial advantage , killing Artybius . However , the defection of two contingents to the Persians crippled their cause , they were routed and Onesilus was killed . The revolt in Cyprus was thus crushed and the Ionians sailed home . Hellespont and Propontis . The Persian forces in Asia Minor seem to have been reorganised in 497 BC , with three of Darius ' s sons - in - law , Daurises , Hymaees , and Otanes , taking charge of three armies . Herodotus suggests that these generals divided up the rebellious lands between themselves and then set out to attack their respective areas . Daurises , who seems to have had the largest army , initially took his army to the Hellespont . There , he systematically besieged and took the cities of Dardanus , Abydos , Percote , Lampsacus and Paesus , each in a single day according to Herodotus . However , when he heard that the Carians were revolting , he moved his army southwards to attempt to crush this new rebellion . This places the timing of the Carian revolt to early 497 BC . Hymaees went to the Propontis and took the city of Cius . After Daurises moves his forces towards Caria , Hymaees marched towards the Hellespont and captured many of the Aeolian cities as well as some of the cities in the Troad . However , he then fell ill and died , ending his campaign . Meanwhile , Otanes , together with Artaphernes , campaigned in Ionia ( see below ) . Caria . Battle of the Marsyas . Hearing that the Carians had rebelled , Daurises led his army south into Caria . The Carians gathered at the " White Pillars " , on the Marsyas River ( the modern Çine ) , a tributary of the Maeander . Pixodorus , a relative of the king of Cilicia , suggested that the Carians should cross the river and fight with it at their backs , so as to prevent retreat and thus make them fight more bravely . This idea was rejected and the Carians made the Persians cross the river to fight them . The ensuing battle was , according to Herodotus , a long affair , with the Carians fighting obstinately before eventually succumbing to the weight of Persian numbers . Herodotus suggests that 10 , 000 Carians and 2 , 000 Persians died in the battle . Battle of Labraunda . The survivors of Marsyas fell back to a sacred grove of Zeus at Labraunda and deliberated whether to surrender to the Persians or to flee Asia altogether . However , while deliberating , they were joined by a Milesian army , and with these reinforcements resolved instead to carry on fighting . The Persians then attacked the army at Labraunda , and inflicted an even heavier defeat , with the Milesians suffering particularly bad casualties . Battle of Pedasus . After the double victory over the Carians , Daurises began the task of reducing the Carian strongholds . The Carians resolved to fight on , and decided to lay an ambush for Daurises on the road through Pedasus . Herodotus implies that this occurred more or less directly after Labraunda , but it has also been suggested that Pedasus occurred the following year ( 496 BC ) , giving the Carians time to regroup . The Persians arrived at Pedasus during the night , and the ambush was sprung to great effect . The Persian army was annihilated and Daurises and the other Persian commanders were slain . The disaster at Pedasus seems to have created a stalemate in the land campaign , and there was apparently little further campaigning in 496 BC and 495 BC . Ionia . The third Persian army , under the command of Otanes and Artaphernes , attacked Ionia and Aeolia . They re - took Clazomenae and Cyme , probably in 497 BC , but then seem to have been less active in 496 BC and 495 BC , probably as a result of the calamity in Caria . At the height of the Persian counter - offensive , Aristagoras , sensing the untenability of his position , decided to abandon his responsibilities as leader of Miletus and of the revolt . He left Miletus with all the members of his faction who would accompany him , and went to the part of Thrace that Darius had granted to Histiaeus after the campaign of 513 BC . Herodotus , who evidently has a rather negative view of him , suggests that Aristagoras simply lost his nerve and fled . Some modern historians have suggested that he went to Thrace to exploit the greater natural resources of the region , and thus support the revolt . Others have suggested that finding himself at the centre of an internal conflict in Miletus , he chose to go into exile rather than exacerbate the situation . In Thrace , he took control of the city that Histiaeus had founded , Myrcinus ( site of the later Amphipolis ) , and started campaigning against the local Thracian population . However , during one campaign , probably in either 497 BC or 496 BC , he was killed by the Thracians . Aristagoras was the one man who might have been able to provide the revolt with a sense of purpose , but after his death the revolt was left effectively leaderless . Shortly after this , Histiaeus was released from his duties in Susa by Darius and sent to Ionia . He had persuaded Darius to let him travel to Ionia by promising to make the Ionians end their revolt . However , Herodotus leaves us in no doubt that his real aim was simply to escape his quasi - captivity in Persia . When he arrived in Sardis , Artaphernes directly accused him of fomenting the rebellion with Aristagoras : " I will tell you , Histiaeus , the truth of this business : it was you who stitched this shoe , and Aristagoras who put it on . " Histiaeus fled that night to Chios and eventually made his way back to Miletus . However , having just got rid of one tyrant , the Milesians were in no mood to receive Histiaeus back . He therefore went to Mytilene in Lesbos and persuaded the Lesbians to give him eight triremes . He set sail for Byzantium with all those who would follow him . There he established himself , seizing all ships that attempted to sail through the Bosporus , unless they agreed to serve him . End of the revolt ( 494 – 493 BC ) . Battle of Lade . By the sixth year of the revolt ( 494 BC ) , the Persian forces had regrouped . The available land forces were gathered into one army , and were accompanied by a fleet supplied by the re - subjugated Cypriots , along with Egyptians , Cilicians and Phoenicians . The Persians headed directly to Miletus , paying little attention to other strongholds , presumably intending to tackle the revolt at its epicentre . The Median general Datis , an expert on Greek affairs , was certainly dispatched to Ionia by Darius at this time . It is therefore possible that he was in overall command of this Persian offensive . Hearing of the approach of this force , the Ionians met at the Panionium , and decided not to attempt to fight on land , leaving the Milesians to defend their walls . Instead , they opted to gather every ship they could and make for the island of Lade , off the coast of Miletus , in order to " fight for Miletus at sea " . The Ionians were joined by the Aeolian islanders from Lesbos , and altogether they had 353 triremes . According to Herodotus , the Persian commanders were concerned that they would not be able to defeat the Ionian fleet and , therefore , would not be able to take Miletus . So they sent the exiled Ionian tyrants to Lade , where each tried to persuade his fellow citizens to desert to the Persians . This approach was initially unsuccessful , but in the week - long delay before the battle , divisions arose in the Ionian camp . These divisions led to the Samians secretly agreeing to the terms offered by the Persians , but remained with the other Ionians for the time being . Soon after , the Persian fleet moved to attack the Ionians , who sailed out to meet them . However , as the two sides neared each other , the Samians sailed away back to Samos , as they had agreed with the Persians . The Lesbians , seeing their neighbours in the battle - line sail away , promptly fled as well , causing the rest of the Ionian line to dissolve . The Chians , together with a small number of ships from other cities , stubbornly remained and fought the Persians , but most of the Ionians fled to their cities . The Chians fought valiantly , at one point breaking the Persian line and capturing many ships , but sustaining many losses of their own ; eventually the remaining Chian ships sailed away , thereby ending the battle . Fall of Miletus . With the defeat of the Ionian fleet , the revolt was effectively over . Miletus was closely invested , the Persians " mining the walls and using every device against it , until they utterly captured it " . According to Herodotus , most of the men were killed , and the women and children were enslaved . Archaeological evidence partially substantiates this , showing widespread signs of destruction , and abandonment of much of the city in the aftermath of Lade . However , some Milesians did remain in ( or quickly returned to ) Miletus , though the city would never recapture its former greatness . Miletus was thus notionally " left empty of Milesians " ; the Persians took the city and coastal land for themselves , and gave the rest of the Milesian territory to Carians from Pedasus . The captive Milesians were brought before Darius in Susa , who settled them on the coast of the Persian Gulf , near the mouth of the Tigris . Many Samians were appalled by the actions of their generals at Lade , and resolved to emigrate before their old tyrant , Aeaces of Samos , returned to rule them . They accepted an invitation from the people of Zancle to settle on the coast of Sicily , and took with them the Milesians who had managed to escape from the Persians . Samos itself was spared from destruction by the Persians because of the Samian defection at Lade . Most of Caria now surrendered to the Persians , although some strongholds had to be captured through force . Histiaeus ' s campaign ( 493 BC ) . Chios . When Histiaeus heard of the fall of Miletus , he seems to have appointed himself as leader of the resistance against Persia . Setting out from Byzantium with his force of Lesbians , he sailed to Chios . The Chians refused to receive him , so he attacked and destroyed the remnants of the Chian fleet . Crippled by the two defeats at sea , the Chians then acquiesced to Histiaeus ' s leadership . Battle of Malene . Histiaeus now gathered a large force of Ionians and Aeolians and went to besiege Thasos . However , he then received the news that the Persian fleet was setting out from Miletus to attack the rest of Ionia , so he quickly returned to Lesbos . In order to feed his army , he led foraging expeditions to the mainland near Atarneus and Myus . A large Persian force under Harpagus was in the area and eventually intercepted one foraging expedition near Malene . The ensuing battle was hard fought , but was ended by a successful Persian cavalry charge , routing the Greek line . Histiaeus himself surrendered to the Persians , thinking that he would be able to talk himself into a pardon from Darius . However , he was taken to Artaphernes instead , who , fully aware of Histiaeus ' s past treachery , impaled him and then sent his embalmed head to Darius . Final operations ( 493 BC ) . The Persian fleet and army wintered at Miletus , before setting out in 493 BC to finally stamp out the last embers of the revolt . They attacked and captured the islands of Chios , Lesbos and Tenedos . On each , they made a ' human - net ' of troops and swept across the whole island to flush out any hiding rebels . They then moved over to the mainland and captured each of the remaining cities of Ionia , similarly seeking out any remaining rebels . Although the cities of Ionia were undoubtedly harrowed in the aftermath , none seems to have suffered quite the fate of Miletus . Herodotus says that the Persians chose the most handsome boys from each city and castrated them , and chose the most beautiful girls and sent them away to the king ' s harem , and then burnt the temples of the cities . While this is possibly true , Herodotus also probably exaggerates the scale of devastation . In a few years , the cities had more - or - less returned to normal and they were able to equip a large fleet for the second Persian invasion of Greece , just 13 years later . The Persian army then re - conquered the settlements on the Asian side of the Propontis , while the Persian fleet sailed up the European coast of the Hellespont , taking each settlement in turn . With all of Asia Minor now firmly returned to Persian rule , the revolt was finally over . Aftermath . Once the inevitable punishment of the rebels had occurred , the Persians were in the mood for conciliation . Since these regions were now Persian territory again , it made no sense to harm their economies further or to drive the people to further rebellions . Artaphernes thus set out to re - establish a workable relationship with his subjects . He summoned representatives from each Ionian city to Sardis , and told them that henceforth , rather than continually quarrelling and fighting between themselves , disputes would be resolved by arbitration , seemingly by a panel of judges . Furthermore , he re - surveyed the land of each city , and set their tribute level in proportion to its size . Artaphernes had also witnessed just how much the Ionians disliked tyrannies , and began to reconsider his position on the local governance of Ionia . The following year , Mardonius , another son - in - law of Darius , would travel to Ionia and abolish the tyrannies , replacing them with democracies . The peace established by Artaphernes would long be remembered as just and fair . Darius actively encouraged the Persian nobility of the area to participate in Greek religious practices , especially those dealing with Apollo . Records from the period indicate that the Persian and Greek nobility began to intermarry , and the children of Persian nobles were given Greek names instead of Persian names . Darius ' conciliatory policies were used as a type of propaganda campaign against the mainland Greeks , so that in 491 BC , when Darius sent heralds throughout Greece demanding submission ( earth and water ) , initially most city - states accepted the offer , Athens and Sparta being the most prominent exceptions . For the Persians , the only unfinished business that remained by the end of 493 BC was to exact punishment on Athens and Eretria for supporting the revolt . The Ionian Revolt had severely threatened the stability of Darius ' s empire , and the states of mainland Greece would continue to threaten that stability unless dealt with . Darius thus began to contemplate the complete conquest of Greece , beginning with the destruction of Athens and Eretria . Therefore , the first Persian invasion of Greece effectively began in the following year , 492 BC , when Mardonius was dispatched ( via Ionia ) to complete the pacification of the land approaches to Greece and push on to Athens and Eretria if possible . Thrace was re - subjugated , having broken loose from Persian rule during the revolts and Macedon compelled to become a vassal of Persia . However , progress was halted by a naval disaster . A second expedition was launched in 490 BC under Datis and Artaphernes , son of the satrap Artaphernes . This amphibious force sailed across the Aegean , subjugating the Cyclades , before arriving off Euboea . Eretria was besieged , captured and destroyed , and the force then moved onto Attica . Landing at the Bay of Marathon , they were met by an Athenian army and defeated in the famous Battle of Marathon , ending the first Persian attempt to subdue Greece . Significance . The Ionian Revolt was primarily of significance as the opening chapter in , and causative agent of the Greco - Persian Wars , which included the two invasions of Greece and the famous battles of Marathon , Thermopylae and Salamis . For the Ionian cities themselves , the revolt ended in failure , and substantial losses , both material and economic . However , Miletus aside , they recovered relatively quickly and prospered under Persian rule for the next forty years . For the Persians , the revolt was significant in drawing them into an extended conflict with the states of Greece which would last for fifty years , over which time they would sustain considerable losses . Militarily , it is difficult to draw too many conclusions from the Ionian Revolt , save for what the Greeks and Persians may ( or may not ) have learnt about each other . Certainly , the Athenians , and Greeks in general , seem to have been impressed by the power of Persian cavalry , with the Greek armies displaying considerable caution during the following campaigns when confronted by the Persian cavalry . Conversely , the Persians seem not to have realised or noticed the potential of the Greek hoplite s as heavy infantry . At the Battle of Marathon , in 490 BC , the Persians took little heed of a primarily hoplitic army , resulting in their defeat . Furthermore , despite the possibility of recruiting heavy infantry from their domains , the Persians began the second invasion of Greece without doing so , and again encountered major problems in the face of Greek armies . It is possible that , given the ease of their victories over the Greeks at Ephesus , and similarly armed forces at the battles of the Marsyas River and Labraunda , the Persians simply disregarded the military value of the hoplite phalanx — to their cost .