The magazine said the caricatures of Ayatollah Ali Khamenei were published in support of protests in Iran sparked by the death in custody of Mahsa Amini, after her arrest for an alleged breach of the country's strict dress code for women.

Tehran's close ally Hezbollah called on "the French government to take firm measures to punish the perpetrators of this act, who attacked what is sacred and flouted the dignity of an entire nation".

Khamenei "is not just the leader of a great country -- he is a religious reference for tens of millions of followers", the Shia movement said in a statement.

Following the publication of the cartoons on Wednesday, Iran summoned the French ambassador to Tehran and closed a research institute affiliated with France's foreign ministry.

In Tehran on Sunday, dozens of Iranians -- most of them religious seminary students -- protested against the publication in front of France's embassy and burned French flags.

Charlie Hebdo, seen by supporters as a champion of freedom of speech and by critics as needlessly provocative, has a style that is controversial even within France.

It has repeatedly caused diplomatic problems abroad for the French government, which has no links with the magazine but faces domestic pressure to defend its right to free speech in an officially secular country.

In 2015 Charlie Hebdo's Paris offices were attacked by gunmen who killed 12 people, following the publication of caricatures depicting the Prophet Mohammed.