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Four truck manufacturers have been fined a record €2.93bn (£2.46bn) by the European Union (EU) for colluding on prices and passing on the costs of emissions-reducing technology.

Daimler, DAF, Iveco, MAN and Volvo/Renault colluded over 14 years to fix prices.

VW-owned MAN avoided a fine as it blew the whistle on the cartel.

Daimler said it regretted these occurrences and took appropriate action some time ago.

"The company has strengthened its internal controls and has intensified its regular and comprehensive employee training with regard to antitrust law and competition law," it said in a statement.

It is the highest penalty the EU has ever imposed on a cartel, and more than double the previous record.

"These truck makers colluded for 14 years on truck pricing and on passing on the costs of compliance with stricter emission rules," the EU statement said.

EU competition commissioner Margrethe Vestager, said the fines were a clear "message to companies that cartels are not accepted."

The truck manufacturers were responsible for nine out of 10 of the medium and heavy trucks produced in Europe, she added.

But the EU said the collusion was not aimed at avoiding or manipulating compliance with the new emission standards, nor did its investigation find any evidence of the use of emissions cheating software.

The investigation found that between 1997 and 2004, meetings were held at senior manager level, sometimes at the margins of trade fairs or other events, to discuss their plans.

This was complemented by phone conversations.

From 2004 onwards, the cartel was organised via the truck producers' German subsidiaries, with participants generally exchanging information by email.

Daimler received the biggest fine at €1.01bn. Sweden's Volvo was fined €670.4m and Iveco, part of Italian truck and tractor maker CNH Industrial, was fined €494.6m.

DAF Trucks, owned by the US group Paccar, received a €752.7m fine.

The four companies admitted wrongdoing in return for a 10% cut in their sanctions.

Scania, another Volkswagen subsidiary, did not settle and will continue to be under investigation.

There are more than 30 million trucks on European roads, which account for around three quarters of inland transport of goods in Europe.