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Boris Johnson has led tributes to the former head of the Royal Marines, who died on Saturday at the age of 54.

Major General Matthew Holmes, who served in Iraq and Afghanistan, was Commandant General Royal Marines from 2019 until April this year.

The Ministry of Defence has launched an investigation into the circumstances of his death.

The defence secretary said Maj Gen Holmes "embodied the spirit of our armed forces".

He had been awarded a CBE in 2019 and was a pallbearer at the Duke of Edinburgh's funeral at Windsor Castle in April.

Mr Johnson tweeted: "I am very saddened to learn of the death of Major General Matt Holmes. My thoughts are with Matt's family and friends at this difficult time, as well as the Royal Marines and Royal Navy who I know will feel this loss keenly."

Gen Holmes, who lived in Hampshire, commanded 42 Commando Royal Marines from 2006 to 2008. He was appointed as a Companion of the Distinguished Service Order for his leadership on operations in Afghanistan in 2007, receiving the honour from the Queen at Buckingham Palace.

Defence Secretary Ben Wallace said: "Across defence, we mourn the tragic loss of one of our own. Major General Matt Holmes embodied the spirit of our armed forces, serving with distinction and rigour for over 30 years.

"I shall always be grateful for Matt's assistance in leading the Marines through the reforms of the Future Commando Force.

"My sincerest condolences lie with Matt's family and those closest to him."

'Served with distinction'

The Ministry of Defence said he had a distinguished career, which also included operational tours in Northern Ireland and Kosovo.

First Sea Lord, Admiral Sir Tony Radakin, said he was a "devoted former Commandant General who had served the nation, the Royal Navy and his beloved Corps with distinction".

"Matt was also a dear friend to many, and a close friend of mine for over 20 years," he said, offering his "deepest sympathy" to his family. "Their loss is the greatest and most painful: we will be there for them now and always."

Commandant General Royal Marines Lieutenant General Rob Magowan said of his predecessor: "We are one family, together, just as we learned, and as we trained, alongside each other at the Commando Training Centre. Once a marine, always a marine."