In and out of prison, he was still able to expand the gang’s influence, seizing control of gold mines in Bolivar state, drug corridors on the Caribbean coast, and clandestine border crossings between Venezuela and Colombia, according to the US state department.
By most accounts, Tren de Aragua spread out of Venezuela when the country entered a humanitarian and economic emergency in 2014 that made crime less profitable, and now is believed to have nodes in eight other countries, including the US.
The group, in part, operates by forming alliances and partnerships with local criminal organisations.
In Ecuador, for example, the gang is believed to work with groups loosely affiliated with Mexico’s Sinaloa cartel, while in Colombia some have alleged that they have worked with members of the left-wing National Liberation Army guerrilla group, or ELN.
Under the Trump administration, US forces have launched dozens of strikes on boats they say are part of a large-scale operation to ferry drugs into the US, including those it claims are linked to Tren de Aragua.
More than 200 people have been killed in strikes since September, according to US media.
But the military has not provided evidence that the attacked boats were carrying drugs or drug smugglers, sparking criticism of the operation and questions around its legality.
Some legal experts have argued that the strikes could violate international law by targeting civilians without offering them due process.
The Trump administration has said the killings are lawful. In a statement to Congress last year, the White House said US President Donald Trump had “determined” that the US was in a formal armed conflict with drug cartels and that crews of drug-running boats were “combatants”.